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shee si ~ f WILLIAM H. DALE
ew i | f SECTIONAL LIBRARY
| ei DIVISION OF MOLLUSKS
VELIGER
A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY,
Berkeley, California
Volume 12
July 1, 1969 to April 1, 1970
Page II
THE VELIGER
Foreword to Volume 12
IT HAS BEEN SAID that everybody talks about the weather but nobody
does anything about it. The same thing could be said about inflation.
We are all painfully aware of its consequences. Yet, we at the Veliger
think that we are doing our share — and perhaps a little more. In spite
of the continually rising cost of paper, printing and postage, over the
years we have effectively lowered our prices. Our first volume could be
obtained at that time for $1.-; it comprised 40 pages and had no illus-
trations. The present volume was sent to our members for their dues of
$8.- — a rise in cost of 800%; but the volume comprises 488 pages and
64 plates, 5 of them in color — an increase in page numbers alone of
over 1200% with the added bonus of the many fine plates.
When the dues and subscription rates for the coming year were set,
very careful consideration was given to the present inflationary trends.
But despite these pressures, the decision was made to keep the dues and
subscription rates at their present low level. This was done not without
the hope that our members and subscribers would assist us in our effort
by helping to increase the number of members and subscribers. We feel
that no one need hesitate to recommend the journal to his conchological
and malacological friends. It also will assist us to keep our overhead
expenses at a minimum if members will pay their dues promptly as well
as advise us in proper time of address changes, thereby making all our
financial resources available for our primary goal — the continued pro-
duction of our journal.
That it has been possible to continue producing our journal under the
ever more stringent financial conditions has been due mainly to the
selfless devotion of a very few individuals. Though they wish to remain
unnamed, your editor would be remiss if he were not expressing his
deep appreciation to each and every one of them.
Sincerely,
Your Editor
Vol. 12; No. 4
Vol. 12; No. 4
TABLE OF CONTENTS
A bibliography of the biological writings of PHP
PEARSALL CARPENTER.
BISGEINIEMV- | COAN) | i seee eeh ns ferret tne esc 222
Additional bathymetric and locality data for some opis-
thobranchs and an octopus from Santa Barbara
County, California.
RICHARD S. LEE & PATRICK BROPHY esses 220
A list of recommended nomenclatural changes for Mac-
FaRLAND’s “Studies of opisthobranchiate mollusks of
the Pacific coast of North America.”
RIGETARDWATIROUT ER: enna ve dol ee. 371
A mechanical shell washer.
JacgueELin N. MILver & Cart L. HUBBS wee 379
An annotated bibliography of references to marine Mol-
lusca from the northern state of Sonora, Mexico.
CS AROMISKOGL UND etd ent can ele cece 427
Anatomical notes on the mactrid bivalve, Raeta plicatella
Lamarck, 1818, with a review of the genus Raeta
and related genera.
Harotp W. Harry
An ecological study of valley-forest gastropods in a mixed
mesophytic situation of northern Kentucky.
BraNnLey A. BRANSON & Donan L. BatcH_............ 333
A new species of gastropod (Fissurellidae, Fissurisepta)
from the eastern North Pacific Ocean.
LASS JM UGHLE, CLONZSceceeee e e 24
A new species of Helminthoglypta from the Mojave
Desert.
WAT TER DB SIVIEE DER titer ochre aia cs Bh 275
A new species of Muricopsis from West Mexico.
GerorceE E. Rapwin & ANTHONY D’ArrItio ............. 351
A note on the chromosome number and interrelationships
in the marine gastropod genus Thais of the United
States Pacific Coast.
MuzamMMiL AHMED & ALBERT K. SPARKS ( oeeccccson 293
A note on the opisthobranchs of Santa Cruz Island, Cali-
fornia.
STEVEN MSONG ti) chb ne ie Sanit SU Oe Od seat rah ets 232
A possible “defense” response in a commensal polychaete.
Ronatp V. Dimock & Joyce G. DimocKk_ ...........0...
A report on the feeding of Dendronotus iris on the antho-
zoan Cerianthus sp. from Monterey Bay, California.
Done RAWOBBERG pate ee eet Ae 383
A revision of the eastern Pacific Ovulidae.
@RAWEORDRING CATE Hee ee een ee ee ee 95
Ascophyllum nodosum: a source of exotic invertebrates
introduced into west coast near-shore marine waters.
RICHARD MU MIBEER 2 eet ee AE 230
A simplified vacuum apparatus for collecting small nudi-
branchs.
ER OEIER MAN BUEAIGNEY 0 cscs ond wate. Memeo eee 142
THE VELIGER
Page III
A supplement to the annotated list of opisthobranchs from
San Luis Obispo County, California.
IRIGEARD/ AW ROLLER eerie tens neat mse eetenat 482
A three-dimensional representation of measurement data.
PAu CHANLEY & W. A. VAN ENGEL sscsecensenens 78
Cadlina modesta: a range extension, with notes on habi-
tat and a color variation.
IANS s BERTS GH) Sige aortee ae anni neg eterna 231
Cargoa cupella, new genus and new species of nudibranch
from Chesapeake Bay and the generic status of
Okenia Menke, Idalia LrucKart, and Idalla
ORSTED.
RosAauie M. Vocet & LEONARD P. SCHULTZ... 388
Concerning the validity of the new species of Paravitrea
proposed by BRANSON & Batcu (1970).
Lanpon T: Ross & LAWRENCE C. ABELE oases 484
Correlation of radula tooth structure and food habits of
three vermivorous species of Conus.
JAMES NYBAKKEN sien eGt seme ontd ooh ose 316
Current paleontological investigations on Cenozoic marine
mollusks of the west coast of North America.
WarreN O. ADDICOTT & SABURO KANNO © erettsnsssessscsee 135
Cuttlebones on the beach at Galveston.
Harowip W. Harry & SELMA FE SNIDER oivssscscsssssnn 89
Cyclostrema miranda BARTSCH, a synonym of Tornus sub-
carinatus (MonTacuU).
DONALD ERG: MOORE hye een rs See ar een ee 169
Cypraeidae of the Red Sea at Massawa, Ethiopia, with a
zoogeographical analysis based on the ScHILDERs’
regional lists.
MiCSonve Le. P RUEBUSH MR a ss inet. cautndad 201
Descriptions of a new genus and eight new species of
eastern Pacific Fissurellidae, with notes on other spe-
cles.
JAMES DEOMGICEAN, «occ .cccetacatarian. tani lech eens 362
Escape response of the sea-anemone Anthopleura nigres-
cens (VERRILL) to its predatory eolid nudibranch
Herviella BABA spec. nov.
RU SIROSIN Mra Sees oe tear a2, ores 74
Escape responses of three Indian molluscs.
ANID VANSELT, 60 cl 0 aed ate es al ee 157
Five new species of Terebra from the eastern Pacific.
Mwita BratcHer & R. D. BURGH tite ident 295
Identification of the feeding types in the genus Conus
LINNAEUS.
(CUE BIGING ee ae esate ean ae oe ae 160
Levels of spontaneous electrical and acetylcholinesterase
activities during aestivation of the Indian apple snail,
Pila globosa.
P Muraut Mowan & P. Murari KrisHna Dass 37
Macoma (Psammacoma) pulleyi, a new clam from Louisi-
ana.
PAUL Ss BOVER ojo siocssccsntsicrsst cn cned ees nee eh ee 40
Page IV
Malacological applications of scanning electron micro-
scopy. I. Introduction and shell surface features.
FAA s SOLE Ms scscentaastsnsriiclassncean aie ae ere een 394
Marine fouling and boring organisms in Monterey Harbor
— II. Second year of investigation.
Ee CHApERTIE ries Oe. See ee eae 182
Mimicry of the gastropod Mitrella carinata by the am-
phipod Pleustes platypa.
JUrESEM@ CRANE: fre SS ae Se ee ee 200
New range for Mopalia hindsi recurvans BARNAWELL,
1960.
GLENN BurRGHARDT & LAURA BURGHARDT oe 229
New species of Panamic marine gastropods.
(AMESMETS Melia Nese ese eee eee 310
Notes on the deep water calliostomas of the Panamic pro-
vince, with descriptions of six new species.
AMES EL MCLEAN: ate ee eee 421
Notes on the egg capsules and larval development of
Conus purpurascens BRODERIP.
PANES WINVBAKIKE Nis, 2208 Dees Binh paar eB ee 480
Notes on the Mitridae of the Eastern Pacific II — The
genus Thala, with the description of a new species.
GALE Gr SPHONI ga, bike at oe neha 2 eae We 84
Notes on the collection of Tritonia festiva (STEARNS, 1873)
from the seas of Japan (Gastropoda: Nudibranchia).
KIKUTARO BABA. ci dea, ee eee ee 132
Observations on the anatomy and biology of two Califor-
nia vermetid gastropods.
MIGHAET| Gi HAD ETER D2 etieetee eis ora een 301
Observations on the reproductive biology of the Kellet’s
whelk, Kelletia kelleti.
RIGHARD!]; ROSENTHAL 3 S22 S00) ee eee 319
Observations on the tentacles of Vaginulus borellianus
Co tosl.
ARISTEO)/RENZONTO encode oe eee 176
Occurrence of a rare squid, Chaunoteuthis mollis APPELL-
6F (family Onychoteuthidae) in the Indian Ocean.
KN ANESIS( 3th ein ne eee 290
Occurrence of the sacoglossan opisthobranch Hermaea
dendritica ALDER & HANcock in New England.
Kerry B. CLarK & DAVID R. FRANZ oresmsssssnseseinaneen 174
On Pseudopythina rugifera (CARPENTER, 1864).
WaLter NarcHioiet ha aioe eee ee 43
Observations on Pervicacia tristis (DEsHayES, 1859) and
a comparison with other toxoglossan gastropods.
WRB RUD MIAN ups hie ce oot ea a 53
Occurrence of the cephalaspid Philine sinuata (Stmmp-
SON ) in southern New England, with a discussion of
the species.
DROP RANZ {EK S GUAR, 1.2.00 200 INE iad Bhd 69
On CEerNoHorsky’s designation of a lectotype for Murex
mancinella LINNAEUS.
Haroip E. Vokes
THE VELIGER
Vol. 12; No. 4
Panamic sites and archaeological mollusks of Lower Cali-
fornia.
LAWRENCE HL. FELDMAN, cieccmcscecneismnsrennnneent 165
Pleistocene symbiosis: pinnotherid crabs in pelecypods
from Cape Blanco, Oregon.
Victor A. ZULLO & DUSTIN D. CHIVERS 0 oun 72
Population characteristics of Protothaca staminea (Con-
RAD) from Mugu Lagoon, California.
Rona_p R. ScomptT & JOHN E. WARME on 193
Range extension of Tjlodina fungina in the Gulf of
California.
James W. McBetu « R. David BOWLUS nese 229
Range extension of Tochuina tetraquetra (PAuas, 1788)
to Hokkaido, North Japan (Gastropoda : Nudibranchia).
KIKUTARO. BABA j jasgeincencithgien tee 134
Range extensions for Acanthodoris hudsoni MacFarLaANnD,
1905, and Onchidoris bilamellata (LinNaEus, 1767).
Gary R. McDONALD: eiscioctiecenniene ee 375
Relative growth patterns of two west coast squid (Gona-
tus fabricii and Gonatopsis borealis).
Larry 'T. SPENGER) ses:ecdtectindinecs Ree eee 171
Remarks on the taxonomic placement of Purpurellus
JoussEaumME, 1880, with the description of a new
species.
WILuIAM K. EMERSON & ANTHONY D’AttTitio —_—«:145
Report on some abnormal chitons from California and
British Columbia.
GLENN BurGHARDT & LAURA BURGHARDT ou... 228
Reproductive cycle of the coot clam, Mulinia lateralis
(Say), in Long Island Sound.
ANTHONY CALABRESE) sencscisncardarhes-ca eee 265
Russian contributions to malacology.
KENNETH J. BOSS. 222 e2 A ae cane 226
Seasonal gonadal changes of adult oviparous oysters in
Tomales Bay, California.
Cart J. Bere, Jricu eee A 27
Spawning notes V. — Acanthina angelica I. OLpRoyp,
1918 and Acanthina lugubris (SowerRBy, 1821).
FAY HENRY, WOLFSON Ges. eenen 375
Supplementary comments on deep water Volutidae from
the South China Sea and South Africa.
HARALD) A. [REHDER "vs. cecchidank destin be ee 415
Technique for extraction and mounting of gastropod rad-
ulae. E
GrorGe, Ey RADWIN( once diecsceinaseeeeenne ae 143
The cowrie species living at Guam.
CRAWEORD! Ni (CATE Goce eee 120
The distribution and ecology of sub-littoral species of
Macoma (Bivalvia) off Moresby Island and in Satel-
lite Channel, near Victoria, British Columbia.
R. M. Dunnit « D. V. Eis
The eastern Pacific cowries.
CRAWFORD. IN. (CATE: 22... cceeentes 28 ae eee, 103
Vol. 12; No. 4
The effect of wave impact on some aspects of the biology
of sea mussels.
Jap cu Ei EVARGER) Geet 2 mere case ojo wane Meant 401
The endodontoid land snail genera Pilsbrycharopa and
Paryphantopsis.
JUIN, SOOTY. Creat cee ch uc art Men ee ee 239
The systematics and some aspects of the ecology of the
genus Dendronotus.
GORDON VAHROBIDETARD: aca Sot fishaseselicctescnedssausimtne 433
The west American species of Murexiella (Gastropoda:
Muricidae), including two new species.
IBMT AEA SAVOKIES! | ccttrnthe termine niet niu tae 325
Three new species of muricacean gastropods from the
eastern Pacific.
WiiuaM K. Emerson & AntTHony D’Attitio 270
Two new Epitoniidae from the Galapagos Islands.
DEVE NAPING YU S EUAN IE | cet actclaccct Oh Peart sgcncssscerisetessseesa 330
Two new species of deepwater bivalves from the Carib-
bean Sea.
EVAR OV DBE MeN OKE Sie ite Mustaine ed adh ean lie al oni 357
Two new species of Galapagan turrid gastropods.
WILuIAM K. EMERSON & GeorGE E. Rapwiw_........ 149
Two new species of the genus Caldukia BuRN & MILLER,
1969 (Mollusca: Gastropoda: Opisthobranchia) from
New Zealand waters.
VET EVAR CMMNITE EER) ites ene oe le bi ua dha 279
Uptake of sea water into the fluid spaces of the proso-
branch gastropod, Acmaea scutum.
IT emE PAVE BBE Rage nite crc ns 0, Part ceetnsuels cel es 417
AUTHOR INDEX
ABELE, LAWRENCE G. see Ross, LANDON T. & —
AppicoTt, WARREN O. & SABURO KANNO
AHMED, MuZAMMIL & ALBERT K. SPARKS
J\SSIGHOH CI ANG) DS, ae IN ae oe Mec re ee
Basa, K1KUTARO
BErc, Cart J., Jr.
IBERTS CHa EVANS) )cnnee nese
BLEAKNEY, J. SHERMAN
Boss, KENNETH J.
Bow us, R. Davin see McBETH, JAMES W. & —
BOYER PAU INS ethene Men ten oe a a eae 40
BRANSON, BranLey A. & Donatp L. BatcH_.......... 333
BRATCHER, TWILA & R. D. BURGE occcssssssssssnsussssssssntsns 295
Bropuy, Patrick see Ler, RicHarp S. & —
Burcu, R. D. see BRaTcHER, TWILA & —
BuRGHARDT, GLENN & LAURA BURGHARDT ........ 228, 229
BurGHARDT, LAuRA see BURGHARDT, GLENN & —
(CATABRESE VAINDHONY 9) tees freon i ua ins Aad L 265
CATES GRAWFORDYNS; Shot. Se OoNLO3S a al20
THE VELIGER
CHANLEY, PAUL & W.A. VAN ENGEL rcsmssssssmssssssnsasssseee 78
Curvers, Dustin D. see ZULLO, Victor A. & —
CiarK, KERRY B. & DAVID R. FRANZ rvsssssssssssssssssneen 174
Crark, K. see FRANZ, D. R. & —
(OLOVAING , VERO SSI VG ce eee cect reer peer renee crmreteecerats 222
Cowan, Ian McT__.............. 24
Crane, Jutes M., Jr. 200
Dass, P. Murat KrisHNna see Mouan, P. Murai & —
D’Attitio, ANTHONY see EMERSON, WILLIAM K. & —
see also: RapDwin, GrEorcE E. & —
Dimock, Joyce G. see Dimock, Ronatp V.
Dimock, RONALD V. & JOYCE G. DIMOCK oeeeecsesnsine 65
ID) einRpacses IS, IML CE LDS WE OVS) ert eromeemnpermeemrenoreremmee 207
DU SHANE, SELELEN set sisstafeoscicescsasissvasectisiatantictewnssssmtaanots 330
Exus, D. V. see DUNNILL, R. M. & —
EMERSON, W. K. & A. D?ATTILIO oivcescsssussssssnsinnee 145, 270
EMERSON, WILLIAM K. & GEorGE E. RADWIN esmeese 149
IBELDMANRICAWREN CE sbi) ee eesreenne nn nnn ee see 165
omnes PaRUEBUSH eee eae 201
BRAN ZI Rese KCI CEARK), ; coer ni ee ee 69
Haverue, E. C.
HaprFietp, MicHaet G.
LARGER Je Rete ater em meres Nestea. es at CAE 401
PARRY TARO ED. Wi crises ies Ue ee Ee ae ea 1
Harry, HAROLD W. & SELMA E SNIDER oovssssssssessssssssnseeee 89
Husss, Cart L. see MILter, JAcQUELIN N.
Kanno, SABuRO see ApDICOTT, WARREN O. & —
LEE, RICHARD S. & PATRICK BROPHY ( osscssssssssssssussssns 220
1 GY o.Y Ge OD Date etre eet
EONG SO TEVEN Po) |. ciseteteece teeta saree accra eareines
McBetH, James W. & R. D. Bowtus
McDona_p, Gary R............
IMGIERAIN SDJ AMES#EN: cha cnren meee canaane
MILLER, JACQUELIN N. & Cart L. Husss
Mier, MicHaet C............
Mitter, RicHarp L. .......
Miter, WALTER B.
Mouan, P Murat & P. Murai KrisHNa Dass ..... 37
Moore, Donatp R.
NARCHI, WALTER ..ececos 43
INESISHKESNe) cecsicesecnnsann 290
NYBAKKEN, JAMES
Rapwin, GrorcE E.
see also: EMERSON, WILLIAM K. & —
143
Rapwin, GrEorGE E. & ANTHONY D’ATTILIO ocecsecscs 351
IREHDERWEVARALD IA cossetiead deccssane ueccsne enn 415
IRENZONIGP ARIS TRON pester ee ie
RoBILLIARD, GorDon A.
IROLEER RICHARD) Ramer tele
RosENTHAL, RicHarD J.
Rosin, R.
IRUDMANEA Wile es ee ie SEE ee ee een a 53
Page VI
THE VELIGER
RuesusuH, L. P. see Fon, T. C. & —
SCHMDDT, RONALD R. & JOHN E. WARME eesti 193
ScHuLTz, Leonarp P. see RosALIE M. VocEL & —
SKOGLUND,; CAROL 11h tea cee ee eee Meeeelinoneree 427
SNDDER, SELMA F see Harry, Harotp W. & —
SOBEM: CALAN fk: cee tet a, 1) (enum elas 239, 394
SPHON, (GALEN Gi), 22 ee. eee eee ee ea 84
Sparks, ALBERT K. see AHMED, MUZAMMIL & —
SPENCER CARRY Ay ee Cee 171
Van ENGEL, W. A. see CHANLEY, PAUL & —
Vokes sEinaiiy He ten hee ccna ee 325
VoGcEL, Rosauiz M. & LEonarp P. SCHULTZ. .............. 388
VOKES; HAROLD) Bier Boa. eee ie ae 357, 368
WarME, JoHN E. see ScummpT, Ronatp R. & —
WEBBER HEHE ih AUR er ee Oe a ene an 417
WoBBER}: Dony. RY hhh. ga a Oe 383
WOLFSON) PAY HENR Yarn ou eee wee ee 375
ZuLLO, Victor A. & DUSTIN D. CHIVERS (ovine Wi2
Vol. 12; No. 4
VELIGER
THE
A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.
Berkeley, California
VOLUME 12 July 1, 1969 NUMBER I
ConTENTS
Anatomical Notes on the Mactrid Bivalve, Raeta plicatella Lamarck, 1818, with
A Review of the Genus Raeta and Related Genera. (20 Text figures)
Haro.p W. Harry
A New Species of Gastropod (Bissrelndae: Rasurieniayy far the Basen North
Pacific Ocean. (3 Text figures)
Tan McT. Cowan
Seasonal Gonadal Changes of Adult @uiesraue Gystersh in Glomale: Bay (Californie:
(Plates 1 to 3; 1 Text figure)
Cart J. Bere, Jr.
Levels of Spontaneous Electrical and Aceurleholinestsrase INctivities durae Nestiva-
tion of the Indian Apple Snail, Pila globosa. (1 Text vee
P Murari Monan « P Murat Krisuna Dass
Macoma (Psammacoma) pulleyi, a New Clam from Tous (2 Text eure)
Paut S. Boyer
On Pseudopythina ueiete (Canesten, 1864) (Bivalvia). Gl Text fegres)
Water NarcHI
Observations on Pervicacia tristis Gieerneee 1859) aaa a Gonpaneen cain oer
Toxoglossan Gastropods. (5 Text figures)
W. B. RupMAN 6 Aue en pniee ae Nana
A Possible “Defense” Reeponseli ina s Commmencal Polychacte (Plate 4)
Ronatp V. Dimock « Joyce G. Dimock
Occurrence of the Cephalaspid Philine shed Greco it in Souler New enelandl
with a Discussion of the Species. (8 Text figures)
D. R. Franz « K. Ciark
Pleistocene Symbiosis: Pinnotherid Gens in Weelenypods) fara cone Bianco! orear
(Plate 5)
Victor A. ZuLLo « Dustin D. CHIVERS
[Continued on Inside Front Cover]
7 a ra pre ee
Distributed free to Members of the California Malacozoological Society, Inc.
Subscriptions (by Volume only) payable in advance to Calif: Malacozool. Soc., Inc.
( gut 24 1969
Volume 12: $18. Domestic; $19.- in the Americas; 19.50 in all other Foreign Countries
Single copies this issue $12.-. Postage extra.
Send subscription orders to Mrs. JEAN M. Carte, 12719 San Vicente Boulevard,
Los Angeles, California 90049. Address all other correspondence to Dr. R. Stouer, Editor,
Department of Zoology, University of California, Berkeley, California 94720
Second Class Postage paid at Berkeley, California
R A R ES
-
ConTENTs — Continued
Escape Response of the Sea-Anemone Anthopleura nigrescens (VERRILL) to its
Predatory Eolid Nudibranch Herviella Baza spec. nov.
Rev ROSINSS ee =), SESE! ee Heer amen Al
A Three-Dimensional Resretentaron of ‘Measurement Data (4 Text figures)
PauL CHANLEY & W. A. Van ENGEL ee ‘ dey Bent eer th nether 7S
Notes on the Mitridae of the Eastern Pacific Ir - The Gann Thala, with the
Description of a New Species. a 6; 2 Text ga I nee
Gate G. SPHON. : : : a) Soa ee Od
Cuttlebones on the Beach at Galveston! (3 Text figures)
Haroitp W. Harry & SELMA FE SNIDER eee Suanaro aes!)
A Revision of the Eastern Pacific Ovulidae. (Pate 7 to 10; et Maps)
CrawForpD N. Cate. . . 2 het Es
The Eastern Pacific Cowries. . eiates) It to 15; 3 Maps)
Crawrorp N. Cate . . Sf cs ee sent epee EOS
The Cowrie Species Living at Guan (Molluses : (Grsttopoday, (Plates 16 to
25; 1 Map)
CRAwrForD NiiGATEs <7 iGer" otis Bee aia ote ee ey an ee nee a eT
INKS) CZINDINIS) co 9 0 0 Baga wo kG See gaat Vora. of os. he IGA
Notes on the Collection of Tritonia festiva iin 1873) from the Seas
of Japan (Gastropoda : Nudibranchia) (1 Text figure)
KIkuTARO BABA
Range Extension of Tochuina tetraquetra (PaLias, 1788) to Hokkaido,
North Japan (Gastropoda : Nudibranchia). KrxuTarO BABA
Current Paleontologic Investigations on Cenozoic Marine Mollusks of the
West Coast of North America. WarreN O. AppICoTT &
SABURO KaNNO
METHODS & TECHNIQUES... . aon ss(/1P)
A Simplified Vacuum Apparatus for Collectine Ae Nae
(1 Text figure) J. SHERMAN BLEAKNEY
Technique for Extraction and Mounting of Gastropod Radulae.
GrorcE E. RaDWIN
Note: The various taxa above species are indicated by the use of different type styles
as shown by the following examples, and by increasing indentation.
ORDER, Suborder, DIVISION, Subdivision, SECTION,
SUPERFAMILY, Famity, Subfamily, Genus, (Subgenus)
New Taxa
Vol. 12; No. 1
THE VELIGER
Division Of Molluske
sctionall Library
Page |
Anatomical Notes on the Mactrid Bivalve,
Raeta plicatella LAMaRcK, 1818,
with
A Review of the Genus Raeta and Related Genera
HAROLD W. HARRY
Texas A&M Marine Laboratory, Galveston, Texas 77550 and Rice University, Houston, Texas 77001
(20 Text figures)
TABLE OF CONTENTS
Mina itech Ur Cte TD es secsesccesces ase ast ieeas vas saasteevsesdvasseesveatsvsesvsssesectontieosestee? I
Part I. Raeta plicatella
a. Ecology
Ip e hia Vl on terete a setacceemterttortnrstntoneccommtionmneste ize 2
c. Anatomy
d. Anatomical Discussion
Part II. Systematics
a. Genus Anatina
b. Genus Raeia s. s.
c. Subgenus Raetina __.......
d. Subgenus Raetella _......
e. Systematic Discussion
Weiteratune Cited) ee ee ede ans QI
INTRODUCTION
SEVENTY YEARS AGO, DALL (1898, p. 907) wrote of this
species:
“Notwithstanding the fact that dead valves of this shell are
found in windrows on the beaches at some points of the south-
ern coast, the character of the soft parts is unknown, and I
shall be very greatly obliged to any one who can furnish me with
a specimen of the animal in spirits in order that its systematic
position may be positively settled.”
Apparently nothing has yet been reported on its ana-
tomy. It may, therefore, be worth recording notes which I
made on a single intact live specimen, and fragments of
two others, for this species has several unusual structural
characters which heighten interest about its enigmatic
ecology and systematic relationships.
Raeta plicatella is the type of its genus. I have therefore
included a review of this genus and related ones. Since
nothing is known of the anatomies of any of the species
of the other genera, conclusions on the systematics of this
group must be based on characters of the shell. Descrip-
tion of the shell of R. plicatella is deferred to the system-
atic part of the paper.
I am indebted to Mr. Pat Burke, Mr. Donald Harper,
and Mr. Ronald Tomas for assistance in field work, and
to the staff of the Mollusk Department of the United
States National Museum for their generous hospitality
while I studied the National Collection.
PART I. Raeta plicatella (Lamarck, 1818)
Ecology:
Single, fresh valves are often abundant on the beaches
along the Gulf of Mexico in Louisiana and Texas. Very
rarely shells have a bored hole, probably made by Poli-
nices duplicata Say, 1822. Joined valves of Raeta plica-
tella are rare and live specimens even more so. I have
found only two shells containing the flesh on the beach.
Both were at the edge of the surf, and both had been
picked open on the central part of the disk by sea gulls.
Birds may account for the dearth of live specimens, even
in the winter when rough surf often tosses up live snails
and bivalves whose shells afford them more protection
Page 2
(Anadara, Noctia, Dosinia, Tellina alternata Say, 1822,
Busycon, Polinices and others).
Raeta plicatella seems to be a species of the outer part
of the surf zone, a region difficult to sample. The mollusks
which live there are chiefly known by being cast up on
the beach by rough surf, for they are not regularly found
in the inner part of the surf zone frequented by swimmers,
and several of them are not taken by dredging in waters
seaward of this zone. At Galveston, the outer surf zone
is only a few hundred yards wide, and from 2 to 3
fathoms depth. It is so close to shore that boats large
enough for trawling and dredging cannot often be used
there. In more than 60 dredged samples from 3 to 18
fathoms off Galveston, single, small juveniles of R. plica-
tella occurred in two samples at 3 fathoms, and single
fragments of larger shells were found in two samples of
5 fathoms. Not even identifiable pieces of shell occurred
in any of the others.
I had never seen shell fragments or juveniles of this
species in the Galveston Bay complex, until the single
intact specimen was found which furnished most of the
anatomical data of this report. It was dredged from 2
fathoms, 14 October 1967, at the east end of the intra-
coastal canal cutting Pelican Island, Lower Galveston
Bay.
Occasional live specimens of mollusks living in the
outer surf zone are found in Galveston Bay. These are
usually juveniles, or rarely small adults. Their scarcity
and size suggest that their larvae have invaded this area,
which is adjacent and connected by water to that in which
they normally live, but that they are not able to complete
their life cycle and establish perpetuating populations in
the bays. Examples of such adventitious species found in
the more saline parts of the bays, but with established
populations only along the Gulf shore are Dosinia discus
Reeve, 1850, Tellina alternata Say, 1822, Anadara brasili-
ana, Periploma inequalis C.B.Apams, 1842, Noetia
ponderosa Say, 1822, Pitar texasiana Dati, 1889, and
Donax variabilis Say, 1822.
Behavior:
In the laboratory, the live specimen was lethargic. It
did not move about in a dish of seawater, and the foot
was not seen extended beyond the shell. The shell gaped
slightly and continuously during the several days of ob-
servation. The siphons were not withdrawn, even when
the specimen was handled.
Several colonies of the stalked ciliate protozoan, Car-
chestum, were attached to the siphons near their tips,
and also found on the periostracum between the margins
of the valves along the postdorsal slope of the shell.
A small colony of an encrusting bryozoan was on the
THE VELIGER
Vol. 12; No. 1
periostracum of the fused mantle margin near the pedal
opening. The surface of the shell was clean and had no
extraneous organisms. The presence of Carchesium and
the bryozoan suggest this animal did not burrow, but was
on its side on the substrate. This inference is further
supported by the nature of the foot (see below).
External Anatomy:
The siphons (slightly contracted in Figure 1) are about
2 as long as the shell, and joined throughout their length.
There is a slight groove between the two, on the sides.
Figure i
Diagram of the external anatomy of Raeta plicatella from a live,
partly retracted specimen. The left valve and mantle have been
removed.
aa -—anterior adductor muscle _ap — anterior pedal retractor muscle
f — foot g—gill Ip — labial palp pa-— posterior adductor
po — pedal opening in mantle margin __ pr — posterior pedal retractor
r—rectum sm-—siphonal membrane sp -—siphonal pocket
wl-—right wall lamella of waste canal v— ventricle of heart
The diameter of the excurrent siphon is about 4 that
of the incurrent one. The opening of the excurrent siphon
has a thin, transparent, conical flange extending from it,
with a small opening at its tip. There are no papillae
around the excurrent siphonal tip. The incurrent opening
has no valve flange, but numerous short papillae are
along its margin. Two papillae of this series, between the
two siphonal openings, are larger than the rest. Every 10
minutes or so the siphons contracted rapidly, but only
for a short distance, and the feces were expelled through
the excurrent opening.
A white, wavy seam on the top and bottom of the
siphons evidently represents the junction of the two sides
of the periostracum. This cuticular sheet extends to the
Vol. 12; No. 1
THE VELIGER
Page 3
tips of the siphons, lines the siphonal retraction cavity
of the mantle, and all of the exposed mantle margin. It
is very thin, and scarcely evident over the outer shell
surface.
The color of the living tissue was varied. There was no
gross evidence of hemoglobin, the blood being colorless
and the nerve ganglia white. The ovary was white, the
liver the usual olive brown, the kidney was faint purplish
brown. The labial palps and gills were faint orange yellow,
and the foot was dingy white. The rest of the flesh was
without notable color.
The mantle is very thin, and colorless over the disk of
the shell. The disk part is attached in small round spots,
which are not the origin of muscle cells, but seem to be
epithelial. The spots of attachment are 1 to 2mm in
diameter, and widely scattered.
The kidney extends into the lobes of the mantle, ex-
tending downward well below the free margin of the
gills (dashed line, Figure 2). This part of the mantle is
thickened.
The margins of the mantle lobes are fused to each other
along the dorsal, anterior and ventral margins of the
shell, and the attachment is thick and muscular. At the
antero-ventral margin there is a small opening in the
x
Oy
as
SC pb
Figure 2
Visceral mass and structures in the mantle at the hind end
of the body.
ap — anterior pedal retractor muscle
ec — excurrent chamber of mantle cavity
i-intestine hp-liver k-—kidney
pb — posterior end of gills
pg—pedal ganglion = pr—posterior pedal retractor muscle
rms-— retractor muscles of siphons _ sc — style sac
s—septum separating mantle chambers sm -—siphonal membrane
sO—sense organ ssg — suspensory septum of gills st — stomach
v-—ventricle of heart vg -— visceral ganglion
cg — cerebral ganglion
ga—gill axis
ov — ovary
marginal fusion, the pedal opening (Figure 1). This
is surrounded on its inner margin by a thin, narrow velum,
which may represent the inner mantle margin lamella.
The latter is not evident as a distinct entity elsewhere.
No “fourth pallial aperture”, below the siphonal pocket,
was evident (see YoncE, 1948). At the hind angle of the
shell, there is a large opening into the deep, blind siphonal
pocket (Figure 1). |
In front of the siphonal pocket, the siphonal retractor
muscles form a flat, triangular mass in each mantle lobe,
with the apex pointing forward. These muscles are small
bundles, not forming a continuous sheet, but with spaces
between them about half as wide as each bundle (Figure
2, rms). The origin of the siphonal retractor muscles is
along the pallial sinus scar of the shell. They converge
toward and extend into the siphons as longitudinal re-
tractors.
A thin, transparent diaphragm of mantle epithelium
extends forward horizontally from between the inner
openings of the siphons. This septum is a continuation of
the diaphragm formed by the gill attachments, and with
the latter it completely separates the incurrent and excur-
rent chambers of the mantle cavity (Figure 2, s).
There is a single lamella of mantle extending along
both sides and the top of the incurrent siphonal opening,
but not across its bottom (Figure 2, sm). This is the
siphonal membrane (Kettoce, 1915). A pair of lamellae
of moderate width extend forward from the ends of the
siphonal membrane, one attached to each mantle lobe,
to a point below the apex of the pallial sinus scar (Fig-
ure 1, wl). These are the wall lamellae of the waste canal
(KELLOocG, op. cit.). These three lamellae have intrinsic
musculature, which on contraction decreases the height
of the lamellae. They probably serve to guide pseudofeces
into the incurrent siphon.
The fused edge of the two mantle lobes ventrally is
very thick (Figure 3), and filled chiefly with mantle
margin muscles. Some of these pass from the position of
the pallial line to the free, or exposed mantle margin, and
thus pull the mantle margin upwards, away from the mar-
gin of the shell. But the bulk of the muscles pass trans-
versely, forming a mantle margin adductor muscle in this
area, as in certain pholads, such as Cyrtopleura costata.
There are a few strands of longitudinal muscles in this
fused mass, particularly on the upper surface.
The periostracum covering the free, ventral surface of
the fused mantle margin has a white streak medially. Just
above this streak are the two minute grooves (Figure 3,
pg) in which the periostracum is formed, between what is
homologous to the outer and middle mantle margin la-
mellae of other bivalves, which do not have fused mantle
lobes in this region.
Page 4
The anterior adductor muscle (Figure 1, aa) is very
elongate and narrow, extending along the anterior curve
of the shell from the front end of the pedal opening to
the front end of the hinge plate. It is merely a broadening
of the mantle margin adductor muscle, and not a separate
entity. The hind adductor muscle is reniform, and small
for the size of the shell. Both muscles were opaque and
white, with no gross evidence of “quick” and “catch”
parts differentiated.
WC wl
Figure 3
Diagram of transverse section of ventral margin of Raeta plicatella,
about one fourth the length from the hind end.
m— mantle lobe
rm — retractor muscle of mantle margin
tm — transverse muscle of mantle margin
wl — wall lamella
Im — longitudinal muscle of mantle margin
peg — periostracal groove
sh — shell
wc — waste canal
The foot is atrophied to a small, elongated, wrinkled
mass, flattened laterally and rounded at the free end. The
free margin is acutely keeled, but without any groove.
Probably the foot is never extended beyond the shell mar-
gin in specimens of this size (44mm long). It is some-
what muscular, the circular muscles extending only a
little way up on the front margin of the visceral mass,
but not up the hind margin of it. Anterior and posterior
pedal retractor muscles each form a thin bundle along
the front and hind margin, respectively, of the visceral
mass (Figure 2, ap,pr). At their top, each muscle divides,
the front one originating on the shell valves on each
side and slightly above the mouth, and thus just above
the anterior adductor muscle.. The hind pedal retractor
has the rectum passing between its two short branches.
THE VELIGER
Vol. 12; No. 1
It originates on the shell valves just above the posterior
adductor muscle scars.
The visceral mass has an exceptionally thin wall, with
little musculature. Probably this is a consequence of the
atrophied foot. The covering is so thin that the internal
organs of the visceral mass are easily seen through it.
The gills have two demibranchs on each side. The
ascending lamellae of all four demibranchs are attached
to the mantle at about the level of the gill axes. The
gill axes form almost a straight line running obliquely
from a point a little way in front of the siphonal inner
openings, forward and upward to a point just below and
slightly in front of the umbos of the shell. The anterior
halves of the attachment of the two gills are separated by
the visceral mass, but behind it they are so joined that
the excurrent chamber of the mantle cavity is completely
separated from the incurrent one. Behind the visceral
mass, the ascending lamellae of the medial demibranchs
are attached to each other along their upper margins. The
gill axes are here each suspended by a thin vertical
septum, so that the excurrent chamber is divided into
three compartments, two lateral and a medial, between
the hind margin of the visceral mass and the lower end
of the hind adductor muscle (Figure 2, ssg). Behind
that point there is only a single tubular excurrent cham-
ber, limited below by the horizontal septum of the mantle,
laterally by the mantle proper, above by the lower surface
of the kidney and the posterior adductor muscle, and
behind the latter, an additional short horizontal septum
of the mantle.
The outer demibranchs are not as wide as the inner,
and do not extend as far forward. They are semilunar in
shape, with short filaments at both ends, increasing grad-
ually and regularly in length to the middle of the demi-
branch. The medial demibranchs extend a considerable
distance forward, and actually begin well in front of the
hind end of the attachment of the labial palps. The
gills are eulamellibranchian, pleated, and all demibranchs
have a groove along their free margin.
The labial palps are large, subtriangular, with convex
anterior margin, concave posterior margins, and rounded
ventral angle (Figure 1). They are attached along a
straight line to the visceral mass by their dorsal margin.
The two free sides of this triangle have a narrow strip
lacking the numerous vertical ridges which cover the
opposing surfaces of the labial palps. These ridges extend
almost to the mouth; the medial parts of the palps are
narrow, with simple free margins.
Only a few major features of the internal anatomy
could be worked out. The two liver lobes, right and left,
are rather small, and fill the periphery of the upper, ante-
rior part of the visceral mass. The gonad, an ovary in the
Vol. 12; No. 1
one complete specimen available, fills the posterior half
of the visceral mass peripherally, pushing dorsally over
the stomach and under the pericardium. The antero-
ventral part of the visceral mass has a large coiled mass
of intestine (Figure 2).
The esophagus is short, opening into a large oval
stomach. The interior of the stomach is much complicated
with ridges and folds, details of which were not worked
out. Most of the interior was covered by a loose lining of
cuticular material, the gastric shield. A sorting area, of
several oblique ridges on the left side, was not covered by
the shield.
Each liver lobe consists of a thin-walled, branched
tube. The liver follicles, glandular in appearance, are not
branched, but clustered about the tips of the smallest
branches of the hepatic ducts. Each follicle is about
0.1 mm in diameter.
Postero-ventrally, the stomach receives the style sac on
the left, and the slightly smaller intestine opens beside it
on the right. The style sac and intestine are completely
separated, the former plunging down to the tip of the
visceral mass and turning forward for a short distance,
ending blindly. A well-formed crystalline style was pres-
ent, after the clam had spent several days alive in the
laboratory, and even after it had been opened for 2 days,
maintained in the refrigerator at 4° C in sea water.
The intestine is thin-walled, light brown, and evidently
without musculature. A dark brown fecal rope was passed
along by ciliary activity. The intestine is unusually long,
about several times the length of the shell. It is mostly
coiled in the lower front part of the visceral mass, in a
part of the hemocoel free from liver and gonad. Two
parts of the intestine are recognizable on the basis of
size, position and internal structure. The first part (duo-
denum) is imbedded in the ovary on the right of the
style sac, and passes forward near the lower end of the
visceral mass; it also forms the first few coils. It is about
twice the diameter of the second part, the jejuno-ileal
intestine. The latter makes up most of the coils, three of
which loop symmetrically from side to side peripheral to
the lower end of the liver; it then passes into the ovarian
mass well ventrally, turns upward behind the style sac and
enters the front end of the pericardium, beyond which point
it may be called the rectum. After leaving the latter it
passes between the two kidney lobes, over the outer sur-
face of the posterior adductor muscle and opens into
the hind part of the suprabranchial chamber of the
mantle cavity, just in front of the excurrent siphon.
I could not determine the relative lengths of the large
and small intestine, nor their junction. Perhaps it is a
gradual transition. The large (duodenal) intestine has a
huge lamella, or typhlosole, in it, much thicker than the
THE VELIGER
Page 5
intestinal wall. At intervals of a few millimeters, this has
short branches at right angles, but only along one side.
These resemble buttresses. I could find no typhlosole in
the small intestine, but perhaps one is present as a mere
strip of cilia.
Feces:
The pseudofeces, consisting of particulate material as-
sembled in the incurrent mantle cavity and rejected
through the incurrent siphon, are poorly consoltdated
ropes of sediment, slightly larger than a millimeter in
diameter and a few centimeters long.
True feces are of two types, which may be called ropes
and pellets (Figure 4). The pellet type was seen in the
Figure 4
Feces. Rope type on right, an unopened pellet left above, and
a pellet cut transversely on left, below.
rectum of the damaged specimens from the beach, and
the rope type in the intestine of the intact specimen
found in the bay. This specimen voided both types in the
laboratory, and from that material Figure 4 was drawn.
The rope type of feces evidently passes through the
intestine with little rearranging of the materials involved.
It consists of a single strand, 80 to 100mm long, with seg-
ments of dark tan material alternating with segments of
dark olive color. The lengths of the two segment types
are not constant. Probably the olive colored material is
derived from the liver, and the tan material is of undi-
gested particles which entered the mouth. These were not
Page 6
studied in detail. The packaged configuration of the rope,
bent back upon itself several times as shown in Figure 4,
probably occurs as the rope is defecated into the supra-
branchial mantle chamber. This package has been passed
from the animal, as had numerous pellets.
The pellet feces were probably in the specimen when
captured. They are constant in diameter, about 0.69 mm,
and slightly variable in length, about 0.96 to 1.23mm
long, with longer pellets being more abundant. Each pellet
is elaborately constructed. The pellets are smooth, regular
cylinders with abruptly truncated ends, one being slightly
convex, the other slightly concave. The surface is uni-
formly light tan colored, evidently of fine particles held
compactly together with mucus. There is a single line,
slightly incised, which begins in the middle of one end,
makes about one complete turn on the lateral surface,
and ends at the middle of the other end of the pellet.
Breaking a pellet open gently reveals a central cavity
which seems not to extend quite to the ends. Its diameter
is about 4 that of the pellet, and.it is reniform in cross
section. This cavity is loosely filled with granules of vary-
ing sizes, all larger than those of the outer part. The
outer part of the pellet has about five layers, probably of
only one continuous sheet, wrapped around the central
cavity. The innermost layer is wider and less firmly con-
solidated than the others. The layers bend toward the
concave side of the central cavity, and the spiral line on
the outside seems to correspond to this also.
At higher magnification, the particles of the outer shell
of the pellet all seem to be inorganic, angular granules,
rarely as large as 3u. No Brownian movement could be
seen in a squash preparation in sea water, possibly be-
cause the mucous matrix was too viscous. The granules
of the central cavity are irregular spheres, from 10 to 14u
in diameter. These are hyaline, golden brown and bright
in reflected light. Also present were a few glassy angular
flakes, colorless and polygonal, with sharp edges and
angles. These are about 0.30 mm in maximum dimension.
I was unable to find any organismal remains in the pellets,
such as diatoms, even at 970 * magnification.
The pericardium is a large, spacious, cuboidal sac
located below the umbos (Figure 2). The ventricle of the
heart surrounds the rectum, which transverses the peri-
cardium lengthwise dorsally. The ventricle has two small
projections antero-dorsally. The auricles are thin-walled
triangular sacs diverted ventro-laterally. They nowhere
touch each other. A light brown color on their exterior
surface suggests a pericardial gland is present, but it does
not extend onto the pericardial wall. There are two
minute reno-pericardial openings on the post-ventral sur-
face of the pericardial sac.
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Vol. 12; No. 1
The kidney consists of two huge sacs, between the
pericardium and the posterior adductor muscle. The pos-
terior pedal retractor muscle passes between these sacs.
Each sac is elongate, but bent upon itself, so that there
is a dorsal moiety, beginning near the pericardium and
passing backward to the adductor muscle, then turning
forward ventrally to form the ventral moiety, which ends
again at the pericardium. The large extensions of the
kidneys into the mantle are projections from the upper
moiety of each sac. The reno-pericardial passage is at the
front end of the lower moiety of each sac, and a little
farther back each lower moiety has a renopore opening
into the excurrent mantle chamber, just in front of the
post-pedal retractor muscle. The interiors of the kidney
sacs are spacious; there are no trabeculae, but a smooth
wall with large cilia, and numerous small holes opening
into it (Figure 5). These are the ends of short narrow
Figure 5
On the left, a part of the wall of the kidney, showing follicles
extending from its outer surface and pores by which they open on
the inside. Cells are shown in only one follicle. On the right,
a single kidney cell from inside a kidney follicle.
ducts of the kidney follicles. Each follicle is branched
two or three times; they are thin-walled, transparent sacs
(about 0.3mm diameter whose cavities are almost oc-
cluded by the bulbous ends of the excretory cells. These
cells are part of the follicular wall, but they extend in-
ward by narrow, short stalks. The free end of each cell is
spherical, having a large vacuole which holds a light
golden fluid. The fluid does not seem to mix with sea
water. Each vacuole also has a spherical mass of golden
brown, opaque granules.
The members of all three pairs of ganglia of the
nervous system are fused, so that there are no commissures
(Figure 2). The connectives between ganglionic pairs
were not traced. The cerebral ganglia are just above the
mouth. They are intermediate in size between the other
Vol. 12; No. 1
two pairs. The pedal ganglia are 0.19mm long, and
located at the front end of the base of the foot. The
visceral ganglia are 0.3 mm long, and on the roof of the
excurrent mantle chamber below the kidneys, but well in
front of the posterior adductor muscle.
There are two prominent sense organs on the mantle,
one on each side of the upper part of the inner end of
the incurrent siphon, just in front of the median accessory
mantle fold. These may be osphradia. Each is connected by
a large nerve to the visceral ganglion on its side. They are
elongate oval masses, about 0.1 mm wide and 0.3 mm long,
and as high as they are wide.
The ovary of the intact specimen was a large, dendriti-
cally branched multifolliculate organ, the follicles being
slightly less than 0.1 mm diameter, containing thousands
Figure 6
Tape worm larva from: cyst on intestine. Terminal sucker everted
in complete specimen, retracted in the smaller figure.
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Page 7
of eggs which appeared to be about ripe. The eggs are
white, about 44, diameter, with a large germinal vesicle
27. diameter containing a nucleolus 13 diameter.
There seems to be a single gonopore, opening into the
front of the pericardial cavity just to the left of the
rectum. If so, the eggs evidently must pass through the
pericardium and kidney to reach the exterior.
Parasite:
A small cyst containing tapeworm larvae was attached
to the outer surface of the intestine just below the stom-
ach. It was white, and at first mistaken for a piece of the
ovary. The cyst was thin-walled, elongate, with an irregu-
lar surface. It contained several dozen larvae, each coiled
to form a sphere. Released from the cyst into sea water,
the larvae squirmed with peristaltic movement and everted
the terminal sucker. They attached with the lateral suckers
(bothridia) and moved along a glass surface by alternately
attaching and releasing them.
The larva (Figure 6) is fusiform, about 54y long. It
is colorless, and the body is filled with small hyaline
granules. The hind end tapers acutely, and the front end
is rounded, with a deep, cup-shaped, muscular terminal
sucker. This can be extended forward, or withdrawn to
the level of the bothrydia. There are four elongate oval
bothrydia, with small bases, attached symmetrically around
the scolex. The cavity of each bothrydium is divided by
two transverse partitions into three chambers (areolae) of
equal size. No spines were seen at 100 x magnification,
nor could any internal anatomical details be made out.
Anatomical Discussion:
Most notable features of the anatomy of Raeta plica-
tella are the extension of the kidney into the mantle, the
thin wall of the visceral mass, the atrophied foot, and
the enormously long intestine. The abnormal ecological
occurrence of the specimen on which most of this ana-
tomical account is based might cast doubt upon some of
the data on habits and anatomy. There is indeed an in-
trinsic conflict between the two: the long siphons and
atrophied foot suggest this is a deep burrower which does
not move about much in later life. An atrophied foot in
late life of bivalves is probably more common than real-
ized. Among species of the Galveston area I have seen
such in Periploma orbicularis Guppy, 1878 and Diplo-
thyra smithit Tryon, 1862, neither closely related to each
other nor to the Mactridae. But of the anatomical char-
acters, I suspect only the small size of this specimen is
abnormal in terms of other features; its presence above
the substrate, as suggested by the epizoa growing on it,
may indeed be unusual.
Page 8
The diversity in soft anatomy of the Mactridae ap-
pears to be larger than it is within most families of
bivalves. I will not here try to summarize all of the liter-
ature on this subject, which, though extensive and scat-
tered, constitutes a mere beginning in comparison to the
number of species whose anatomies are unknown. A well-
developed foot is present in all species previously studied,
and the atrophied foot of Raeta plicatella is unique. Fu-
sion of the mantle margin behind the pedal opening
varies. In some species, the mantle lobes are completely
free between the anterior adductor and the siphonal pock-
et. In others, the lobes are extensively held together along
the posterior half of their margins by a fusion of a
cuticular extension of the periostracum (Lutraria, fide
Yonce, 1948). Dati (1898a) found free mantle lobes
with papillae in Resania, but in Zenatia he says they are
fused along their hind half, although he does not say
whether the fusion is cuticular or represents cellular con-
tinuity. In Raeta the fusion is definitely cellular, and this
genus therefore represents a third condition of the mantle
margin.
A “fourth pallial opening” which is a small hole be-
tween the mantle edges just below the siphonal pocket,
is present in some Mactridae and other families, according
to Ke.ioce (1915) and Yonce (1948), but such an open-
ing is absent in Raeta.
The wall lamellae of the waste canal, as these structures
were termed by Kettoce (1915), have been given only
passing attention in the literature except by that author
and YoncE (1948), who termed them “mantle folds”.
Dati (1898a) described similar folds in the New Zealand
genera Resania and Zenatia, but called them “sensory
lamellae”.
The siphonal lamellae and wall lamellae of the waste
canal may be structures limited to the Mactridae, al-
though YoncE says that folds similar to the wall lamellae,
but probably differing in function, occur in the Tellina-
cean genera Scrobicularia, Abra and Macoma. A siphonal
lamella forms a complete ring in the venerid Saxidomus
as described and figured by Kettoce, but whether this is
homologous to the semicircular lamella of the incurrent
siphon of the mactrids is unknown. Among mactrids,
the siphonal lamella is said to be absent in Lutraria
(YonceE, 1948), and both that and the wall lamellae are
absent in Spisula planulata Conrad, 1837, according to
KELLOGG.
Whether or not gills are pleated or smooth, without
vertical folds seems to be a character which varies at the
generic or specific level. RmEWoop (1903, pp. 237 ff.)
found flat, homorhabdic (i.e., not-pleated) gills in four
species of Mactridae on which he reported. Whether the
pleated gills of Raeta plicatella have heterorhabdic fila-
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Vol. 12; No. 1
ments at the grooves between the pleats was not deter-
mined.
The persistence of a crystalline style in bivalves which
are not feeding, or at least while they are in the labora-
tory for some time with minimal food, is a phenomenon
which seems to be characteristic of most marine ones I
have studied. This contrast with the oysters and unionids,
which seem to resorb the style quickly when not feeding,
and reform it in the- presence of food.
The opening of the gonad into the pericardium, so that
the gametes transverse the kidney lumen and emerge
through the renopore is a point which bears checking in
further studies. PELSENEER (1911, p. 100) thinks such
a condition is primitive, and limited to some protobranchs,
Anomiidae and Pectinidae. From his reservations on the
matter, it is evident that the gonopore is difficult to
detect in bivalves.
Athough Moore (1931) called attention to the varied
form of fecal pellets in marine mollusks, and suggested
that careful study might yield valuable data on phylo-
geny and food habits, nothing further seems to have been
published on the subject. Moore did not describe the
pellets of any mactrid bivalves. The feces of Raeta
plicatella are markedly different from any he described in
Pectinidae, Mytilidae, Tellinidae and Semelidae. The elab-
orate pellets of R. plicatella are very similar to those of
the pholad C'yrtopleura costata LinNAEUs, 1758, but
differ in minor details (unpublished data).
The presence of an osphradium in bivalves, homologous
to that of snails, is a moot point. Only occasionally have
such structures described above as an osphradium been
noted. PELSENEER (1911, p. 112) calls these “sensory
organ of the inhalent siphon”, and cites them in the
Mactridae.
The parasite found is evidently a larval tapeworm of
the order Tetraphyllidea, the adults of which live in the
intestine of elasmobranch fishes. Larvae of this type are
called “Scolex pleuronectes Mit ver, 1788” or “Scolex
polymorphus Rupotput, 1819”, binomials which do not
have exact taxonomic significance. As Suip.ey (in SoutH-
WELL, 1925, pp. ix-x) points out, “It is extremely diffi-
cult to identify the larval form of a cestode with its
parents. It is a wise tapeworm that knows its own father”.
Apparently, the structures of the scolex in these larvae
may vary considerably at different stages (ages) (SouTH-
WELL, 1925, pp. 138 - 140). The one found in this study
might be assigned to the family Phyllobothriidae, in
lacking hooks on the scolex, yet the general form of the
bothrydia is more like that of the other family in this
order, Onchobothriidae, which, however, has no hooks
in the scolex. Although most of these larval tetraphyl-
lideans have been found in teleost fish, Hyman (1951, p.
Vol. 12; No. 1
354) notes that they have been found in invertebrates of
several groups, including mollusks, and she figures one
(J. c., figure 131 D) from the foot of a clam.
Part II. SYSTEMATICS
Review of Raeta and Related Genera:
Anatina ScHUMACHER, 1817
Raeta Gray, 1853 has sometimes been considered a dis-
tinct genus, a position adopted here. Other writers have
considered it indistinguishable from, or only a subgenus of,
Anatina SCHUMACHER, 1817, or its junior synonym, Labi-
osa Miter, 1832. The nomenclatural problem of Anatina
versus Labiosa has been recently explored by KEEN (1961,
q. v. for relevant bibliography), who gave good reasons
for accepting the name Anatina as valid in the Mactridae.
Moreover, she has shown that the type species of Anatina
SCHUMACHER, 1817 is a western Atlantic species, Mactra
anatina SPENGLER, 1802. This has been cited in much of
the literature as Anatina or Labiosa lineata Say, 1822,
but is now correctly identified as Anatina anatina (SPENG-
LER, 1802).
Despite KEEN’s study, the problem of whether to ac-
cept Anatina SCHUMACHER or Labiosa MULLER has not
been put to rest. OLsson (1961, p. 332) accepts Labiosa,
as does Vokes (1967, p. 274), both apparently on the
assumption that a use of the generic name Anatina by
Bosc in 1816 is a validly proposed name, also a junior
synonym of Laternula “BotTEN” Ropine, 1798. The
latter is a relative of the anomalodesmacean genus Peri-
ploma. According to that line of reasoning, Anatina would
therefore not be available to use in the Mactridae. The
point of contention seems to revolve on what Bosc (1816,
p. 492) said in defining Anatina; his description is here
quoted:
“ANATINE, Anatina. Genre de coquilles bivalves
établi par Lamarck aux dépens des Solens, sous la
consideration d’une petite lame saillante 4 chaque
valve en dedans ow s’attache le ligament.
“Le Solen canard sert de type 4 ce genre, que est
fort voisin des corbules et des Rupicoles. (B)”.
If we consider, as Keen has correctly done, the type
cited by Bosc to be vernacular and therefore invalid, one
might still argue that this is a genus validly described but
without species. Whereas it may be entirely possible to
conceive of a genus without species in the realm of pure
thought, as may be done in modern logic, such a concept
is absurd when applied to real biological entities.
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Page 9
Anatina anatina is a relatively rare shell on the beaches
from New Jersey to Brazil. At Galveston it has not been
found in the bays, but only as shells on the sandy beach
and in a few dredged samples off shore. Apparently it
does not occur in the Antilles.
The differences between the shell of Anatina anatina
and the species from the West Coast of the Americas have
been discussed by Keen (1961), who pointed out that
the West Coast species should be called Anatina cyprinus
Woop, 1828. I could find only a single valve of the latter
species in the National Collection. It is from Topolobampo,
Sinaloa, Mexico. Otsson (1961) has given a descrip-
tion and further distribution records of this species under
the name Anatina anatina. As far as I can determine, the
two species from the coasts of the Americas are the only
ones which should be put in the genus Anatina ScHuUMA-
CHER.
The synonymy of the genus Anatina and the two spe-
cies included in it will not be repeated here, since KEEN
(1961) has provided a meticulous account of these earlier
in this journal, and I have nothing to add. The anatomies
of the soft parts of these species is unknown, but it is
worth pointing out that many recent authors (e. g., YONGE,
1948) refer to the anatomy of the “‘Anatinacea” when
they are referring to the anatomy of the Anomalodesma-
cea of modern classification, i.e., the Periplomatidae,
Lyonsiidae, Pandoridae and related groups. This confusion
results from the fact that PELSENEER (1911 and earlier
works), who paid scant attention to nomenclature, used
the term Anatinacea based on Anatina as a genus in
Periplomatidae. PELSENEER wrote most of what is known
on the structure of the bizarre Anomalodesmacea, and
most recent anatomists have unfortunately accepted his
competency in nomenclature no less than in anatomy.
The shell of the type species of Anatina SCHUMACHER
has been studied in more detail than has been previously
reported, and a description of it will be useful for com-
parison with Raeta and related genera.
Anatina anatina (SPENGLER, 1802)
(Figures 7 to 10)
Shell very thin, fragile, white, translucent, elongate ovoid,
the front end somewhat more sharply rounded than the
hind. Size moderate (65 mm long, 43 mm high, 28 mm
wide). Equivalve, and slightly inequilateral, the umbos
being just behind the midpoint of the length. Umbos
touching, minute and distinctly prosogyrous. The hind end
of the shell gapes. The umbonal fourth of the shell has
faint concentric corrugations, parallel with the growth
striae, forming ridges and grooves on the inner surface.
Page 10
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Vol. 12; No. 1
The lower three fourths of the shell is smooth. A lunule
is poorly defined by a shallow sulcus extending from the
umbos in a broad arch about half the length of the
Figure 7
Anatina anatina, dorsal view. Galveston, Texas, length 43 mm.
antero-dorsal margin. This becomes wider toward its ante-
rior end. On the medial side of the lunular sulcus is an
opaque line not structurally differentiated on the outer
surface. This is the line of attachment of the anterior
hinge plate. The post-dorsal margin is about two-thirds as
long as the antero-dorsal one. It is strongly turned later-
ally, enhancing the posterior gape of the shell. A low rib,
scarcely higher than wide, extends from the umbo to the
post-ventral part of the shell: this may be a corcelet la-
mella. In front of the rib, and separated from it by a
space about half as wide as the space behind the rib, is
a line defining an abrupt change in shell texture. This may
be called the periostracal line. Along a narrow but poorly
defined strip in front of the periostracal line vermiculate
sculpture is evident. This is made up of minute wrinkles
in the shell surface, joining and branching and mostly
Figure 8
Anatina anatina. Galveston, Texas, length 33 mm
directed radially. They are not present over the smooth
part of the disk in larger shells, but are uniformly distrib-
uted, although very faint, on all the adumbonal corru-
gated region. In fresh shells, the surface of the thin,
persistent, faint tan periostracum is silky in reflected
light. But behind the periostracal line, the periostracum
is lusterless, and less persistent. There are only coarse
growth lines in the shell behind the line, and a moderately
thick layer of opaque chalky material is deposited there
between the periostracum and ostracum,
The disk of the shell is distinctly swollen along a line
from the umbo toward the antero-ventral angle. This
gives the umbos the appearance of being directed back-
ward, whereas in reality they are turned forward at their
tips.
Figure 9
Interior of same valve as shown in Figure 8.
The pallial line is narrow, located close to the ventral
edge of the valve, with slightly irregular dorsal margin.
The anterior adductor scar is tear shaped, located direct-
ly on the pallial line; it has a pallial line extending dorsal-
ly from it to the front end of the hinge plate. Just above
the anterior adductor scar the dorsal pallial line has a
slight swelling, probably representing the anterior pedal
retractor scar. The pallial sinus is large, rounded in front.
It extends forward to a point below the umbos. Its lower
side is well above the pallial line, not confluent with it.
The posterior adductor scar is subtriangular, and the
posterior pedal retractor scar constitutes a small con-
stricted area confluent with the adductor at the dorsal tip
of the latter. The corcelet ridge is evident internally as a
shallow groove.
Vol. 12; No. 1
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Page 11
The hinge plate is slightly thicker than the valve. The
front and hind halves extend along the front and hind
dorsal margins for about half the length of the latter.
The front half of the hinge plate is thus slightly longer
than the other; this part of the hinge plate is strongly
oblique to the valve surface, so that there is a deep space
between the two. Its free margin is deeply excavated in a
gentle curve, so that the trough between the hinge plate
and valve is minimized. The hind half of the hinge plate
is almost flat and horizontal, with its free margin almost
straight. The resilifer is a triangular pit below the umbo,
with its lower margin projecting as a rounded curve
below the hinge plate.
Figure 10
Anatina anatina. Hinge of shell. Upper is right valve. The valves
are of different shells.
In the left valve, the cardinal tooth is moderately
thick, and the sinus between the arms is large and deep.
The sides of both arms are flat. The hind arm is about
half as long as the front one. It projects over the resilifer
as a free point. There is a thin antero-lateral lamella
separating the front arm of the cardinal from the front
hinge plate. A similar low, thin lamella is a little way
behind the resilifer pit, separating it from the hind hinge
plate. These two lamellae reach to the free margin of
the plate, forming large oblique angles.
The attachment of the tensiliar ligament is a semi-
lunar depression on the shell margin, beginning at and
extending behind the umbo. This depression extends over
the tip of the resilifer pit, concealing it. The front half
of the curved (ventral) margin of the ligament pit is
turned medially, and to its lower (front) surface is fused
a small lamella projecting as a minute spine over the
resilifer pit. This “spur” lamella is parallel to the hind
ramus of the cardinal, separated from it by a deep groove
which receives the hind cardinal ramus of the right valve.
In the right hinge the resilifer and ligamental shelf
are much the same as in the left. There is also a single
posterior lamella a little way behind the resilifer, separ-
ating it from the shallow trough of the hind hinge plate.
The cardinal and antero-lateral lamellae are different
from those of the left hinge. The cardinal is chevron
shaped, with the posterior arm straight and about half
as long as the anterior arm; the dorsal surface of the
hind arm is fused to the medially turned edge of the
ligamental plate, and the lower end of this arm extends
free over the cavity of the resilifer; the medial (free)
margins of the two arms have a deep excavation where
they meet at the apex of the chevron. The anterior arm
is sharply curved in its upper half, by being bent forward.
This part is fused with the umbonal part of the antero-
lateral lamella, which projects free from the lower part of
the front arm of the cardinal, as a minute but thick and
consistent structure.
The lateral lamellae of this species are probably homo-
logous to the lateral teeth of other heterodonts, and indeed
of other mactrids such as Spisula and Mactra. But in
Anatina and Raeta they are more like cardinal teeth,
radiating from the umbo, but not paralleling either the
shell margin or the margin of the hinge plate; there is
moreover only a single front and hind lateral lamella in
both the right and left valves, whereas in Mactra, Spisula
and some other genera, there is a single one before and
behind in the left valve which fit between a pair at each
place in the right valve. Also, in the latter genera, the
laterals are parallel to the shell margin and, of course, to
the margin of the hinge plate. The oblique, single lateral
lamellae of Anatina and Raeta remind one of the laterals
of the Cardiidae, which curve out as low ridges in a
radiating arc from the umbo, on the inner surface of
the shell.
Whatever the soft anatomy of Anatina may disclose,
the characters of its shell seem sufficiently diverse from
Raeta to separate the two generically; the presence of
the periostracal line and the lateral flexing of the post-
dorsal shell margin in Anatina are characters not present
in Raeta. A lunule is defined in Anatzna, but not in Raeta;
the antero-ventral swelling of the disk is less pronounced
Page 12
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in Anatina; the corrugations are limited to the umbonal
third of the shell in Anatina, but extend to the ventral
margin in Raeta; the corcelet ridge is pronounced in
Anatina, but its presence is indicated by a mere cessation
of the corrugations near the post-dorsal margin, with no
radial ridge per se in Raeta. In the right valve of Raeta,
the anterior lamella is directly in line with the front ramus
of the cardinal tooth, which is flat, and the ramus and
lamella are separated by a notch (Figure 12). In A. ana-
tina, the front ramus of the cardinal of the right hinge is
bent above, almost as long as the front lamella, which is
beside it and free from it below (Figure 10).
The vermiculate texture is common to both Anatina and
Raeta. The periostracal chalk deposit, so pronounced in
Anatina, may have become a lost character in Raeta s.s.,
but it is present in Raetina.
Raeta Gray, 1853
The genus Raeta Gray, 1853 (Ann. Mag. Nat. Hist., Ser.
2, vol. 11, p. 43) was briefly described and typed by
original designation and monotypy, “R. campechensis’’,
which isa junior synonym of Lutraria plicatella LaMARCK,
1818.
Raeta plicatella (Lamarck, 1818)
(Figures 11 to 13)
Lutraria plicatella Lamarck, 1818, Hist. Nat. Anim. sans Vert.
5: 470. Not figured; refers to GMELIN p. 3257, and CHEM-
nitz Conch. Cab. 6, tab. 23, fig. 231 with a question mark.
Type locality, “Probablement l’Océan indien.”
Lutraria canaliculata Say, 1822, Journ. Acad. Nat. Sci. Phila.
2: 311 - 312. Not figured. Type locality, “Maryland and as
far south as East Florida.”
Mactra campechensis Gray, 1825, Ann. of Philos., N.S., 9:
135. Not described or figured; refers to “List. 304 (sic)
f. 141”, evidently Lister, 1770, the second or Huddles-
ford edition, where f. 141 of plate 308 (not 304) is a
creditable figure of this species, with the words, “a sinu
campeche” evidently indicating the type locality as the
Bay of Campeche, Mexico.
Raeta perspicua Hutton, 1873. Cat. Marine Moll. New Zea-
land, p. 65. Not figured. Type locality (erroneously) New
Zealand. 1913, SurzrR, Man. New Zealand Moll., p. 970,
Atlas (1915), plt. 60, fig. 5. Oxtver (1923: 184) notes
that this is a synonym of R. canaliculata Say, incorrectly
attributed to New Zealand:
Labiosa (Raeta) plicatella Lamarck. 1917, Lamy, Journ. de
Conchyliol. Ser.-4,-42: 353 - 354; plt. 7, fig. 6 (photograph
of holotype). 6%
The description and figures cited by Lamarck in de-
scribing Lutraria plicatella were insufficient to recognize
this species, and he furthermore thought it came from the
Indo-Pacific area, but expressed doubt. Lamy (1917)
published a photograph of the holotype of Lamarck’s
species which is unmistakably the species of the western
Atlantic coasts. Meanwhile, it has been extensively cited
in the literature under the trivial name applied by Say,
and in combination with various generic names. HUTTON
redescribed and named this species from a shell which he
attributed to New Zealand, but which had evidently come
from elsewhere, as pointed out by Oxtver. There are
apparently no living species of Raeta or Raetella in New
Zealand.
Figure 11
Raeta plicatella, dorsal view of shell 25 mm long, Galveston Beach.
The hole was probably made by a shore bird.
Shell of moderate size (length 72, height 60, width
40 mm), thin, white, ovoid in side view, equivalve and
gaping slightly behind; subequilateral, the umbos being
slightly closer to the hind end. Umbos small, but swollen
and prominent, touching each other and turned slightly
but distinctly forward. The whole outer surface has prom-
inent concentric ribs, rounded, poorly defined and with
the grooves between slightly wider. These extend through
the shell structure, producing a corrugated inner surface.
In shells longer than 55 mm, the ribs tend to be more
irregular and obliterated along the margin, with the inner
surface smooth there. Fine radial wrinkles, closely spaced,
are present on the ribs. These are most prominent on the
lower third and hind slope of larger shells, where they
extend into the grooves as well, and tend to anastomose.
The wrinkles are part of the outer shell layer. The thin,
light tan periostracum is smooth, closely adherent, but
absent from even slightly worn shells. The front dorsal
margin is gently convex, and continues in an even curve
into the ventral margin, which is more convex. The
hind dorsal margin has about the same curvature as the
front, but it is shorter, slopes more abruptly from the hori-
Vol. 12; No. 1
zontal; it meets the ventral margin in an obtusely rounded
angle. There is no lunule; a narrow corcelet is poorly
defined along the post-dorsal margin, extending to its
post-ventral angle. This is formed merely by the con-
centric ribs turning abruptly umbonad along this line,
becoming much weaker, and lacking the vermiculate
micro ridges present in front of the line.
The disk is greatly inflated from the umbo to the antero-
ventral angle. A wide, shallow trough separates the disk
from the post-dorsal slope, which is somewhat flattened.
The shell margin inside is smooth and sharp, the sur-
face of the interior is subporcellaneous. The pallial line is
thin, but prominent and continuous, with a large sinus
reaching half-way forward and with rounded or usually
acutely pointed tip. Its upper border has a rounded angle
midway its length. The lower border is well separated
from the pallial line proper. The anterior adductor muscle
scar is thin and very elongate, narrowing acutely above.
Figure 12
Right (upper) and left hinges of Raeta plicatella. The tensiliar
ligament is present in the left valve only, but the two halves of the
resilium are present in their respective valves.
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Page 13
The hind one is oval to subquadrate, with a small posterior
pedal retractor scar confluent at its upper end.
The hinge plate is only slightly thicker than the rest of
the shell. The hind half is vertical and flat, and of uniform
width. It extends along the upper two-thirds of the hind
margin. The front part has the shape of an acutely ta-
pered triangle; it is about a third as long as the hind
part. It is attached obliquely to the shell. Directly below
the umbo there is a large triangular resilifer, deeply ex-
cavated and extending to the tip of the umbo, but its
upper third is covered by shell. The base of the triangle
is well below the rest of the hinge plate margin. The lig-
ament is entirely internal, and divided into resilium and
tensilium, well separated. The resilium has the form of
two tapering horns, slightly curved, with their apices in
the covered upper parts of the resilifers, directly below
the umbos, and the faces of their circular bases meeting
in the midline. The faces of the opposed bases are con-
cave, with only the peripheral margins touching. The
concavity extends upward on the medial side of the horns
to the tip of each. The outer, lateral side of each horn is
flattened and about a third again as large as the opposing
faces, so that the two halves of the resilium have the shape
of an hour glass when viewed along the median plane of
the animal.
The tensiliar part of the ligament is elongate lanceo-
late, short, attached at the shell margin beginning just
below the umbos, and with its axis sloping ventrally,
away from the shell margin behind. Only the anterior tip
is visible externally. The tensilium is attached to a hinge
buttress, a lamella which parallels the hind side of the
resilifer triangle, and which extends almost to the free
hinge margin. It is obliquely attached to the hinge plate,
so that its medial surface, to which the tensilium is at-
tached, slopes downward and backward toward the resi-
lifer cavity. Its upper third is attached to the shell margin
completely roofing over the subumbonal tip of the trough
on the hind half of the hinge plate. The lower half of
the tensiliar lamella is acute, and extends over the trough
slightly.
Along the tensiliar lamella, below the tensilium, there
arises an oblique buttress lamella, of the hind ramus of
the cardinal tooth. It completely conceals the upper
fourth of the resilifer cavity, covering the apical end of
the resilium. Along the margin of this lamella which ap-
proaches the cardinal tooth, there is a sharp spine pro-
jecting toward the middle of the base of the resilifer.
A single cardinal tooth in each valve has the shape of
an inverted V, with the rami diverging at slightly less than
a right angle, and the apex directly below the umbo,
separated from it only by the thin hinge margin. The left
cardinal fits into the cavity of the right one. The outer
Page 14
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Vol. 12; No. 1
faces of the left cardinal are smooth and flat, but the
two rami are thick and fused so that the cavity of its
chevron is nearly filled solid, leaving only a vague medial
sulcus on the ventral side. Each cardinal tooth is about
half as high as the width of the hinge margin, and the
hind ramus of each projects over the resilifer.
A deep groove along and above the base of each ramus
of the left cardinal receives the right cardinal. This groove
Figure 13
Upper: Raeta plicatella from Galveston, 63 mm long.
Lower: Raeta undulata from Topolobampo, Mexico, 71 mm long.
is formed behind by the margin of the subtensiliar butt-
ress and its prominent spine, and in front by a lamella
of moderate thickness, normal to the hinge plate and
parallel to the anterior ramus of the left cardinal. This
is the anterior lateral lamella. Its umbonal half is low,
and connected to the inner side of the shell by a small
buttress parallel to the hinge plate; this is the anterior
buttress of the cardinal tooth and it reofs the extreme
tip of the anterior hinge plate trough. The lower half of
the anterior lamella is raised into a quadrate tooth-like
process.
In the right valve the hind ramus of the cardinal tooth
is fused with the spine of the buttress arising below the
tensilium, but not completely, so that it and the spine
appear as a bifid tooth. The front ramus is the same length
as the hind one, but it appears to be twice as long, because
of a quadrate lamella directly below and in line with it.
It is separated from the quadrate lamella by a small but
consistent notch. This quadrate lamella is the antero-lat-
eral lamella. The front ramus of the right cardinal has a
thick lamellar buttress extending from its free margin
outward and upward to the inner surface of the shell,
and roofing the trough of the hinge plate.
In both valves there is a single posterior lateral lamella,
running from the umbo toward the ventral margin. This
lamella is bent upward toward the shell margin, so that its
attachment on the hinge plate is oblique, and poorly
defined in medial view. Evidently these two lamellae are
directly opposite each other in the closed shell, and one
does not insert above the other, since neither crosses the
median plane. But they may be homologous to the poste-
rior lateral teeth of other mactrids.
Raeta (Raeta) undulata (Goutp, 1851)
(Figure 13, lower shell)
Lutraria undulata Gou.p, 1851, Proc. Boston Soc. Nat. Hist.
4: 89. Not figured. Type locality: La Paz, Lower Califor-
nia. 1853, Gouxp, Boston Journ. Nat. Hist. 6: 391; plt. 15,
fig. 7.
Oxproyp (1924) has copied the English translation of
the description by GouLp of this species which he pub-
lished in 1853. She also republished (J. c., plt. 21, fig. 11)
his original figure, an external view of a left valve. His
figure 7 also includes a dorsal view of both valves, showing
a prominent posterior gape. The figure he published
shows very regular corrugations from beak to ventral
margin, but in the larger specimens I have seen, these
ridges tend to be obliterated toward the lower margin,
more so than in Raeta plicatella. Perhaps his figure was
drawn from a juvenile shell.
Vol. 12; No. 1
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Page 15
The shell of Raeta undulata is similar to that of R.
plicatella in the coarse corrugations well covered by ver-
miculations, and in the hinge, which also has an oblique
postero-lateral lamella in each valve. It differs chiefly in
the more rounded, posterior end of the shell, in having
the umbos about midway the length or slightly forward
of that point, instead of well behind, and the corcelet
line along the postero-dorsal shell margin is distinctly
farther from the margin than in the east coast species.
The chalk deposit of the periostracum seems to be absent.
More extensive synonymies are given by Otsson (1961),
all to secondary references however. This species may have
the same ecological distribution as the one on the East
Coast, just beyond the surf zone along sandy shores.
KEEN (1958) notes that “Beach valves are fairly common,
but entire specimens are hard to find, even by dredging”.
(Raetina) Dat, 1898
Raetina Dau, 1895, Proc. Malacol. Soc. London 1: 212, Type
R. indica DALL (nom. nud.).
Raetina Dau, 1898, Trans. Wagner Free Inst. Sci. 3 (4) : 822,
Type by O. D. and M Raeta (Raetina) indica Datx, 1898,
which is a junior synonym of Mactra pellicula REEVE.
This is similar in size and shape to Raeta s.s. The hinge
plate is thick, almost vertical behind, and the groove on
it is shallow and rounded in cross section. Sculpture is
continuous top to bottom, and from front to hind end; it
consists of corrugations evident on the inner surface of
the shell, and the corrugations are in line with the growth
lines, not oblique. Vermiculate texture is present, but very
weak and limited in area.
The main reasons for recognizing this as a distinct sub-
genus are two: (1) the sculpture is more delicate, the
corrugations being finer and more closely spaced than in
the typical subgenus; (2) the posterior lamella, running
obliquely from the umbo across the front end of the hind
hinge plate, is absent in this subgenus. Chalky deposits
in this periostracum, which occur in Anatina, may be
present in Raetina, but seem to be absent in Raeta s. s.
This genus was essentially a nude name as of 1895,
diagnosed only by indication of the type species, which
was not described until 1898.
Raeta (Raetina) pellicula (REEvE, 1854)
(Figures 14 and 15)
Mactra pellicula “DEsuayEs” Reeve, 1854 (May). Conch.
Icon 8: Monograph Mactra, plt. 21, fig. 124. Type locali-
ty: Japan.
Mactra anatinoides Reeve, 1854. Conch. Icon. 8: Monograph
Mactra, plt. 21, fig. 123. Type locality unknown.
Mactra pellicula DesuayeEs, 1855. Proc. Zool. Soc. London 22:
68. Not figured. Type locality: “Japan (col. Cuming) .”
Raeta grayi H. Avams, 1872. Proc. Zool. Soc. London, p13;
pit. 3, fig. 23. Type locality: Borneo.
Raeta abercrombiei Metviti, 1893. (In MEtvitt &« ABER-
cromBIE, 1893) Manchester Lit. & Philos. Soc. Mem. &
Proc. Ser. 4, 7: 32 (nom. nud.) ; MrtviLx, 1893, Ibid., p.
65; plt. 1, fig. 25. Type locality: Bombay, India.
Raeta (Raetina) indica Daur, 1895. Proc. Malacol. Soc. Lon-
don 1: 212 (nom. nud.) .
Raeta (Raetina) indica Dat, 1898. Trans. Wagner Free Inst.
Sci. 3 (4) : 882 - 883, footnote. Not figured. Type locality:
Bombay, India.
Anatina (Raetina) indica Dau, 1925. Proc. U.S. Nat. Mus.
66 (17): 2; plt. 20, fig. 2. Holotype.
Raeta jickelii Sturany, 1905. Nachrichtsbl. d. deutsch, Mala-
kol. Ges. 37t year, pp. 133 - 134; text figs. a, b, and c, p.
133. Type locality: “Massaua” (Massawa, Ethiopia, south-
west coast of the Red Sea).
Raeta magnifica Yokoyama, 1922. Journ. Coll. Sci. Tokyo,
44: 132; plt. 8, figs. 12, 13. Type locality: Fossil, Tega
(Kizaki), Japan.
Raeta fragilis GraBau & Kinc, 1928. Shells of Peitaiho, pp.
190 - 192; plt. 7, fig. 54. Type locality: Peitaiho (Yellow
Sea, China).
This species has a very wide range, from the shores of
the Red Sea, to India, Borneo, China and Japan. It evi-
dently shows some variation in form, particularly in the
Figure 14
Holotype of Raeta (Raetina) indica Dart (= Raeta pellicula).
The dark shading along the postero-dorsal margin indicates peri-
ostracal chalk deposit. Length, 43 mm.
length and acuteness of the rostrum. The short-rostrate
forms are what authors have called Raeta anatinoides
(REEVE) and, judging from the figure, R. grayi H. Apams.
ToMLIn (1924) pointed out that the species names pub-
Page 16
lished by Reeve (1854) in his monograph of Mactra
predate all species named by DESHAYES in 1854 and 1855,
even though REEvE attributes many names to DESHAYES.
For exact date on the several species, see the paper by
Tomun. Elsewhere Tomun (1931) declares R. aber-
crombici identical with R. grayi after comparing the holo-
types of the two species. Although Dati was quite sure
his R. indica was distinct from R. abercrombiei, the only
differences he cited were “form and proportion”. The
holotype of R. indica Dat shows nosignificant differences
from a shell in the National Collection from Japan, which
in turn fits quite well with the concept of R. pellicula
Reeve. No one who has studied the group seems to have
seen more than a few shells from different places, and
the above synonymy can only be tentative.
Figure 15
Interior of same valve as shown in Figure 14.
Like the species of Raetas. s., all records of this species
are from the beaches of continents or the larger conti-
nental islands. It may be absent in the small oceanic is-
lands of the Indo-Pacific, which seem indeed to have
few mactrids of any genus. Probably this species lives at
the outer edge of the surf zone on sandy shores.
The following description is based on the holotype of
Raeta indica Daut, which is a single left valve, USNM
90276, with the words “Bombay” and “Wesleyan Univ.”
on the label.
Length 43, height 28, semidiameter 9 mm. Shell elon-
gate ovoid, front end evenly rounded, hind end drawn
out in an obtusely rounded rostrum. Probably equivalve,
and almost equilateral, the umbos being slightly closer
to the front end. Umbos prominent, touching, turned
slightly forward. Shell very thin, translucent, with numer-
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Vol. 12; No. 1
ous concentric ribs and troughs, which are undulations of
the shell substance affecting both surfaces. The undula-
tions are regular, from umbo to ventral margin, and con-
tinuous, except along the post-dorsal slope. They are
parallel to the growth striae. Color white. No lunule.
Corcelet (or escutcheon?) defined only by rapid dimi-
nution of the ribs along a poorly defined line near and
parallel to the post-dorsal margin. No epidermis is evident,
but there is a chalky white deposit postero-ventrally. A
few faint vermiculations, similar to those of Raeta plica-
tella, are evident only along the post-ventral part of the
valve.
The roundly pointed pallial sinus reaches as far for-
ward as the umbo. Its lower margin is separated from
the pallial line. Muscle scars could not be seen in the
slightly worn holotype (but Datu illustrated them as
typical of Raeta s.s.). The hinge is typical of Raeta,
except that there is no hind radiating lamella between the
resilifer and the groove of the post hinge plate. The com-
parable buttress of the front hinge plate groove is slightly
broken, but it has a thick base. The posterior gape
between the valves must have been small.
I cannot see any specific differences in the holotype and
a pair of matched valves from Japan (Hirase Coll., US
NM 344967, 53 mm long, 41 mm high). The Japanese
specimen is larger, not as acuminately rostrate behind.
There is no chalk deposit on its posterior slope, but a
faint, narrow corcelet ridge seems to be present, slightly
forward of the line medial of which the concentric ribs
become mere growth striae, though pronounced.
The form of both the Indian and Japanese shells is
easily distinguishable from Raeta plicatella and R. undu-
lata, the ribs are more numerous and finer, the posterior
lateral lamella is lacking in both specimens of the oriental
species.
Raetella Datu, 1898
Raetella Dax, 1895. Proc. Malacol. Soc. London 1: 212. Type
by O.D. and M. Raetella tenuis (Hinps MS.) Day
(nom. nud.) .
Raetella Dau, 1898. Trans. Wagner Free Inst. Sci. 6 (4):
882 - 883. Type species by O. D. Raetella tenuis “Hinps”
Dat, 1898, which is a junior synonym of Poromya pul-
chella A. ADAMS & REEvE, 1850.
Raetellops Hae, 1952. Genera of Japanese Shells, Pelecypoda
No. 3: 197, as subgenus of Raeta Gray, 1853. Type by
O.D., Poromya pulchella A. ApaMs & REEvE, 1850.
This name was essentially nude when proposed by Dati
in 1895, being diagnosed only by designation of the type
species, which was not described until 1898.
Vol. 12; No. 1
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Page 17
Shells of this genus are much smaller in size than those
of Raeta, and the ribs of Raetella are proportionately
coarser. They are, moreover, slightly oblique to the lines
of growth. The shells have a peculiar opalescent luster,
which Datu called a “‘pseudo-nacreous surface”. Vermi-
culate texture and chalky periostracal deposit are absent.
Although Dati (1898, p. 882) said the “dorsal areas [are]
well defined”, there is no structural feature on the external
surface defining them; their presence is merely simulated
by the attachment of the hinge plate showing through
the translucent shell. The hinge plate lamellae are very
thin, and the front one is only slightly shorter than the
hind one. Both front and hind parts are almost horizontal,
with the space between them and the adjacent valve
surface very deep. In Raeta and Raetina the posterior half
of the hinge plate is essentially vertical and thick, with
swollen, rounded free margin. The lateral teeth are paral-
lel to the hinge plate margin, not strongly oblique to it,
as in Raeta and Raetina. Moreover, there is a single
hind lateral in the left valve, but two in the right, in the
pattern quite general in Mactridae. But there is only a
single, short antero-lateral in each valve. This genus may
extend to greater depths than do Raeta or Raetina, which,
like most mactrids, are shallow water inhabitants. As
presently known, it seems limited to the eastern coast of
Asia and larger continental islands along it.
Raetella pulchella (A. ADAMS & REEvE, 1850)
(Figures 16 to 19)
Poromya pulchella A. ADAMS & REEVE, 1850. Voy. Samarang.
Zool., Mollusca, p. 83; plt. 23, fig. 1. Type locality: Shores
of Borneo.
Mactra rostralis “DESHAYES” REEvE, 1854. Conch. Icon. 8,
Monogr. Mactra, plt. 21, fig. 119. Type locality: “China
Seas?”
Mactra rostralis DEsHaYEs, 1855. Proc. Zool. Soc. London, p.
69. Not figured. Type locality: Japan.
Raeta yokohamensis Pirssry, 1895. Cat. Marine Moll. Japan
. collected by F Stearns, p. 119; plt. 3, figs. 4, 5. Type
locality: Yokohama, Japan.
Raetella tenuis “Hinps” Datu, 1898. Trans. Wagner Free Inst.
Sci. 3 (4): 883 (footnote). Not figured. Type locality:
Hong Kong Harbor, about 8 feet of water.
Raeta elliptica Yokoyama, 1922. Journ. Coll. Sci. Tokyo 44:
131 - 132; plt. 8, fig. 7. Type locality: Fossil, Tega, Ka-
menari, Kioroshi (Japan). ;
Raetella rostralis has been considered a synonym of R.
pulchella by most authors, as pointed out by Lamy (1917).
On the basis of the original descriptions and illustrations
of the several nominal species, I agree with Hape (1952)
that Raeta yokohamensis and Raeta elliptica should be
added to the synonymy. The specimen on which Dati
based his name Raetella tenuis ““Hinps” also seems con-
specific. Hinps seems never to have named such a species,
and Dau (1898) received the name from P. P. Carpenter,
who evidently took it from a label in the British Museum.
The following description is based on the holotype of
Raetella tenuis “Hinps” Datu. The type lot is USNM
519, and the label has written on it “4-8 ft., mud, in har-
bor, Hong Kong, Stimpson”. The holotype consists of
disjoined valves of one shell; the right valve is only
broken around the ventral and posterior margin, but the
left is fragmented into several large pieces.
Shell small (13 mm long, 10mm high, 3.0 mm semi-
diameter), ovate in profile, evenly rounded in front and
Figure 16
Holotype of Raetella tenuis “Hinps” Dati (= Raetella pulchella) .
Length, 13 mm.
Page 18
below, drawn out in a short but rather acute rostrum
behind. The disk is very inflated, almost hemispherical,
but the inflation is even, without a prominent axis from
umbo to antero-ventral margin. Umbo 5/7 the length
from the front end.
Color white. A thin, closely adhering periostracum is
light tan. The outer surface has a faint opalescent irides-
cence. Sculpture of about 23 concentric ribs, which are
regularly spaced from umbo to ventral margin, but which
do not extend to the antero-dorsal margin nor onto the
rostrum. There is a slight, very broad, shallow sulcus be-
tween the disk and the post-dorsal region, but no demar-
cation of an escutcheon or corcelet. There is no chalky
deposit in the periostracum, and no radiating vermiculate
texture. The concentric ribs are slightly oblique to the
lines of growth, and this is most prominent on the antero-
ventral part of the disk. A lunule is simulated by the
attachment of the anterior hinge plate.
Shell very thin, the ribs forming undulations on the
inside, which is porcellaneous. Attachment scars not evi-
dent in the holotype.
Figure 17
Hinge of holotype of Raetella tenuis “Hinps” Dati
(= Raetella pulchella)
In the right valve a V-shaped cardinal tooth is adjacent
to and points toward the umbo. The two arms are about
equal in length, moderately thick, and with a broad, deep
space between them. A deep cleft separates the medial
part of their junction, at the apex of the V (poorly
shown in Figure 17; see Figure 19). The posterior arm
is separated by a narrow space from the front edge of
the resilifer. Along the base of the anterior ramus there
is a very thin buttress which crosses obliquely the space
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Vol. 12; No. 1
Figure 18
Upper: Raetella pulchella from Japan (USNM 249247) showing
pallial sinus. Lower: Raetella pulchella from Fukura, Awaji, Japan
(USNM 344966) 23 mm long. Note relatively smooth surface and
poorly rostrate hind end.
over the cavity of the anterior hinge plate and attaches to
the shell. A lamella in line with the anterior arm of the
cardinal and also the anterior hinge plate lamella seems
to be broken off, flush with the latter, in the right valve
of the holotype (see Figure 19). The anterior hinge plate
is very thin, with sharp free margin. It is almost horizon-
tal, paralleling the dorsal margin of the shell and at-
taching rather far laterally. The cavity between the front
hinge plate and the shell is thus very deep and narrow.
The front hinge plate lamella is only slightly shorter than
the hind one. The hind hinge plate is not so wide as the
front, and the cavity between it and the shell is not so
deep. No escutcheonal area is defined on the outer sur-
face by its attachment. It is equally thin, with sharp
margin. Along the umbonal fourth this plate is extended
medially slightly, forming a lateral tooth. Above it on the
margin of the shell is another very slight lamella, not as
long as the lower one, forming an upper posterior lateral
tooth. The resilifer is small, triangular, directed down-
ward from the umbo. Tensilium very short, directly under
the umbo, and not separated from the resilifer by any
shelly material except the small ledge on which it sits.
Vol. 12; No. 1
THE VELIGER
Page 19
Hinge of the left valve similar to that of the right.
The two rami of the V-shaped cardinal tooth are thicker,
smaller, and evidently fit inside the right cardinal. Above
the left cardinal and between it and the umbo is a small
but prominent tooth-like process, with an inverted V-
shape, the front ramus of which is very short. The hind
ramus of this little tooth margins the upper part of the
resilifer. A lamellar extension of the anterior hinge plate
seems to be present at the umbonal end but it is badly
broken. There is no dorsal lateral tooth on the shell mar-
gin. A very thin prominent lamella radiates from the
umbo and separates the hind hinge plate from the cardinal
area. Two small lamellae in the front hinge plate cavity
of the left valve do not extend to the median plate of
the shell, and are possibly abnormal.
~~ Cho ee Z
~~ 5 Sy?
Sas Bae
SNS ae eee nS
Ses i LELEF
SWS al =
Figure 19
Hinge of Raetella pulchella from Japan (USNM 249247).
The drawings were made at slightly different angles, causing the
resilifer of the left (lower) valve to appear smaller than that of
right hinge.
The holotype of DaLt’s nominal species is more inflated
over the disk and more rostrate behind than most speci-
mens of the 14 lots in the National Collection from Japan.
It fits the concept of “Mactra” rostralis better than “M.”’
pulchella, but I think it is only an extreme variant of
this highly variable species (cf. Figure 18). Some shells
from Japan are larger (20mm long, 14mm high). The
Japanese shells are slightly gaping behind, and equivalve.
Sculpture on them varies considerably, some shells being
almost smooth, with ribs limited to the ventral part, not
present on the umbo. Most Japanese shells are less rostrate
behind than is the holotype of Da.t’s species; they are
more equilateral, and not as inflated over the disk.
Figure 19 is a drawing of the hinge of a Japanese
specimen, showing the prominent short anterior lateral
tooth, directly on the hinge plate margin. Each tooth
curves outward from the umbo, as shown by the dotted
lines at their bases. The left antero-lateral slips above
the right one when the valves close. The left valve has a
single postero-lateral tooth, which fits between the two
on the right valve. All these postero-laterals are developed
as elongate lamellae directly on the valve margin and
hinge plate margin, and parallel to them.
A large lot from Siam (USNM 477293) has very thin
shells which are non-rostrate, well sculptured, and of a
uniform small size.
SPECIES INCERTAE SEDIS
Raeta meridionalis Tate, 1889
(Figure 20)
Raeta meridionalis Tate, 1889. Trans. Proc. Roy. Soc. South
Austral. 11: 61; plt. 11, fig. 3. Type locality: Aldinga Bay
(South Australia).
?Labiosa meridionalis Tate, 1889. HEpLEy, 1900. Proc. Linn.
Soc. New South Wales 25: 497; plt. 25, figs. 5 - 9. Locali-
ty: Chinaman’s Beach, Middle Harbour (Australia).
Tate’s original description of this species is here quoted,
and Figure 20 (left) is a tracing of his original drawing.
“Shell whitish, very thin, translucent; ovately-suboblong, mod-
erately convex antemedially, posteriorly gaping; umbo sub-
acute, curved forward, situated in the anterior three-sevenths.
“Anterior margin regularly rounded, ventral margin strongly
arched; dorsal line sloping on both sides, more so in front than
behind; posterior side narrowed and depressed, its margin
somewhat squarely truncated and slightly reflected. There is an
ill defined depressed post-dorsal area, but no umbonal ridge.
“Sinus widely rounded, horizontal, reaching to the center,
visible on the exterior.
“Surface marked by fine lines of growth, which become more
conspicuous and finely wrinkled on the post-dorsal area.
“Dimensions: Antero-posterior diameter, 35; umbo-ventral di-
ameter, 30; sectional diameter of left valve, 8 millimeters.
“Locality — one valve collected by Mr. Magarey on the beach
of Aldinga Bay.”
The species described by Tate has the shape of Mactra
anatinoides REEVE, which I have considered a junior syn-
onym of Raeta pellicula. But the sculpture is most un-
usual for a Raeta. It is apparently so thin that the pallial
sinus shows through the shell, yet strong concentric cor-
rugations seem to be lacking, and there is no corcelet line
or ridge along the post-dorsal slope. I have seen no speci-
mens. This species evidently lives near the shore of sandy
Page 20
THE VELIGER
Vol. 12; No. 1
beaches and it may be quite rare. A copy of the original
figure is given by Cotron & Goprrey (1938) and by
ALLEN (1959). Incidentally, Cotron’s (1961, p. 343)
reference to the original description of this species (“TATE,
1887, Trans. and Proc. Roy. Soc. So. Austr. 9: 68 - 69, pl.
5, fig. 4, Montacuta meridionalis’) is evidently in error,
for the description and figure of Montacuta meridionalis
Tate, 1887 clearly refers to a leptonid bivalve, and not to
R. meridionalis (Tate, 1889).
PL. XXV.
7 eS
es / Cae —
rE 2) SSS
Sere ——
LG fy \)
i ee i if
Bile
“
, 4 Y
if [ :
Hai AAv!
nti Wo zat
Wo yy
as Hl 4
9S ye
Figure 20
Left: Tracing of figure 3 of plate 11, TaTE, 1889, the original
drawing of Raeta meridionalis Tate. Right: Figures 5-9 of plate
25 of HEDLEY, 1900, which he also identified as
Raeta meridionalis TATE.
The problem is further confused by the account Hep-
LEY (1900) gave of 14 additional specimens which he
found. Although his shells were reported to be authenti-
cated by Tate, the size, shape and sculpture all suggest
a species of the Raetella pulchella group. The peculiar
drawing of the sculpture also suggests the concentric
undulations are slightly oblique to the lines of growth,
as in Raetella, and not vermiculate textural features char-
acteristic of Raeta and Raetina. HeDLEy’s remarks and
figures are here reproduced (Figure 20, right).
“This species has hitherto been known from a single valve
found on the beach of Aldinga Bay, South Australia. This year
I have taken a whole shell containing part of the animal, and
on another occasion a broken valve on “Chinaman’s Beach”,
Middle Harbour. Prof. Tate, to whom one valve was submitted,
kindly informs me that there is no essential difference between
it and the type of meridionalis. My specimens are smaller,
being 28 mm in length and 21 mm in height. Being perfect,
I have utilized my example to draw the valves in apposition
and other details not obtainable from the single valve hitherto
known.
“Though disagreeing by vermiculate sculpture, the species
seems to me nearer to the subgenus Raetella, DALL, than to any
other division of Labiosa.”
Possibly there are two species along the shore of south-
ern Australia, one described by HeEpxey being similar to
Raetella pulchella, the other to a sub-oval Raeta (Rae-
tina) pellicula. SMrrH (1914, p. 150) cites Cypricia grayi
H. Apvams (which is Raeta grayi H. ApaMs cited above
in the synonymy of Raeta pellicula) from Queensland,
Australia, and notes a reference which I have not seen,
which does the same (HeEptey, 1910, Rep. Austral. Assoc.
Adv. Sci. for 1909, p. 351).
Nick es (1950, p. 209) cites a “Labiosa vitrea Gray”
from the west coast of Africa (“Casamance a la Guinée
frangaise; Congo frangais”), with a brief description and
figure. The general shape is that of Raetella, but the size
is larger (up to 50 mm long). There are regular undula-
tions only on the posterior slope, which is unusual for
this genus. This is possibly Mactra vitrea Gray, 1837
(Charlesworth’s Mag. Nat. Hist., New Ser., vol. 1, p.
372, not figured) described from unknown locality. Lamy
(1917, pp. 273 - 274) placed this in the subgenus Mact-
rinula of Mactra, but did not figure or describe it, nor did
he know the locality of the material which he examined
in the Paris Museum. I have seen no specimens, and
cannot comment on its possible membership in Anatina
or Raeta.
As Lamy (1917) has noted, Raeta lyrata “Hinps Ms.”
H. « A. Apams (1856) is a nomen nudum, and Raeta
bracheon Sturany, 1899 (p. 12; pl. 3, figs. 1-6; Gulf
of Suez, 50 fathoms) belongs elsewhere. Lamy says it is
Leptomya cochlearis Hinps, of the Scrobiculariidae.
The Raeta tenera “DESHAYES” listed by H. & A. ADAMS
possibly refers to “Mactra tenera Desh.” listed as a syn-
onym of Mactra anatinoides, with query, by Reeve. As
Tomutn (1924) implies, no such species seems to have
been described by DEsuay Es.
According to the Zoological Record, Li (1930) de-
scribed a new species, Raeta maxima, but I have been
unable to locate that article.
Vol. 12; No. 1
THE VELIGER
Page 21
SYSTEMATIC DISCUSSION
Much essential information relevant to the phylogeny and
life habits of this group is likely to be gained by further
investigation of their anatomies. There are also characters
of the shell which have not been given adequate atten-
tion. Admittedly, the species of this review were chiefly
considered on the basis of those traditionally grouped to-
gether. Anatomical studies on these and other mactrid
species may well indicate revisions of the classification
quite different from that proposed by Dax (1895, 1898)
and expanded by Lamy (1917). Several examples of
shell characters whose significance is unexplored occurred
to me during the present study. One is that thin-shelled
mactrids generally have corrugated concentric sculpture,
with the ridges affecting both the inner and outer shell
surface, as in Raeta and Raetella, or they have a keel
along the post-dorsal slope as in Anatina, Mactra alata,
and M. fragilis; this keel I have called a corcelet, although
aware that it may not be homologous to structures so
named in other families. Perhaps the corrugations and
keel are devices which strengthen the shell. In some
species both are present, notably Harvella elegans from
the tropical part of the western shores of the Americas.
Mactra iheringi (Dati, 1897), from the east coast of
South America, is very similar to H. elegans in size, shape
and dentition, but it is a much thicker shell, with smooth
external surface and no corrugations or post-dorsal ridge.
That these two represent analogous species on the two
sides of the continent should be given further study.
Incidentally, Harvella elegans seems to have vermicu-
late texture similar to that of Raeta plicatella, and the
distribution of this feature among thin-shelled mactras
with corrugated shells should be studied.
In a rare species of the Indo-Pacific, Mactra (Mactrin-
ula) plicataria Linnaeus, 1767, the front end of the shell
is disproportionately elongate, with the hind end trun-
cated, and a form recalling Donax. This genus has further
developed a thin shell, with corrugate sculpture and ver-
miculate texture very similar to Raeta. The disk, however,
is not exceptionally swollen.
In the group of Anatina, Raeta, and Raetella there is a
more pronounced swelling of the disk, which is not me-
dial, but extends more toward the front end of the shell.
In its extreme form, such as found in Raeta s.s., a shape
recalling that of Cuspidaria results. This is just the op-
posite of the disk swollen along a line running postero-
ventrally such as found in the Mytilidae and Carditidae.
Raetella may be no more closely related to Raeta and
Raetina than any of them are to Harvella or Mactrinula.
The sculpture of Raetella recalls that of the West Ameri-
can Tumbeziconcha, which, however, has the posterior
end of the shell extended, and the obliquity of the ridges
on that end, rather than on the front one.
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1961. South Australian mollusca.
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1851. Descriptions of new shells from California, collected by
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1837. A synoptical catalogue of the species of certain tribes or
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1853. A revision of the genera of some of the families of con-
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1952. | Genera of Japanese shells.
187 - 278
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1873. Catalogue of the marine Mollusca of New Zealand with
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Hyman, Lissy HENRIETTA
1951. The invertebrates: Platyhelminthes and Rhynchocoela,
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Keen, A. Myra
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi + 624 pp.; illus.
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1961. | What is Anatina anatina? The Veliger 4 (1): 9 to
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1915. Ciliary mechanisms of lamellibranchs with descriptions
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Lamarck, JEAN-BartisTe PreRRE ANTOINE DE MONET DE
1818. Histoire naturelle des animaux sans vertébres.
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1917-1918. Revision des Mactridae vivants du Muséum d’Histoire
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Li, C. CHanc
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1893. Descriptions of twenty-five new species of marine shells
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THE VELIGER
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MELVvILL, JAMES Cosmo & ALEXANDER ABERCROMBIE
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Page 23
Page 24
THE VELIGER
Vol. 12; No. 1
A New Species of Gastropod (Fissurellidae, Fissurisepta)
from the Eastern North Pacific Ocean
IAN McT. COWAN
Faculty of Graduate Studies, The University of British Columbia, Vancouver 8, British Columbia, Canada
(3 Text figures)
Fissurisepta SEGUENZA, 1863, Is CUSTOMARILY regarded as
a subgenus of the genus Puncturella. This designation has
been based upon characteristics of the shell, and the living
animal seems never to have been described. Recently the
discovery of a single living specimen in a dredge haul
from 470 fathoms off the west coast of British Columbia
not only extends the known distribution of the subgenus
into the eastern Pacific Ocean but also permits reexamina-
tion of its systematic status on the basis of some soft-parts
anatomy.
The type species, Puncturella (Fissurisepta) papillosa
SEGUENZA, 1863, is known only from the Miocene of
Sicily and adjacent regions. A second species of the sub-
genus, P (F}) rostrata SEGUENZA, 1863, was described from
the same deposits. A third species of Miocene occurrence
was described as P (F) retula Wooprina, 1928. His speci-
mens were from Jamaica.
At almost the same time the subgenus was discovered
as a fossil it was found in the living fauna off Norway
(JEFFREY, 1882) and described as Puncturella (Fissuri-
septa) granulosa JEFFREY, 1882. Subsequent explorations
have yielded specimens from several parts of the Atlantic.
Thus P(E) acuminata Watson, 1833, was taken off North
Culebra Island in the West Indies, and again off Yucatan
and off Cumberland Island, Georgia. Specimens from the
two last localities were described by Dati (1889) as
Fissurisepta triangulata, a name now regarded as a syn-
onym of P (F) acuminata (FarFanTE, 1947). This species
was collected again off the Azores where it provided the
source of yet another apparent synonym microphyma
DAUTZENBERG & FiscHER, 1927 (FARFANTE, 1947).
A second species from the West Atlantic off Georgia
was named by Dati (1927) as Puncturella tenuicula.
CiarKE (1962) has named a 5" species from the Atlantic
as Puncturella (Fissurisepta) agulhasae.
Fissurisepta was unknown in the Pacific until 1951 when
Kuropa named and described the shell details of Punctu-
rella (Fissurisepta) soyoae from Japanese waters. More
recently OkuTANI (1964) has reported a second species,
P. (F) undulata from the western Pacific Ocean.
The bathymetric range of this creature, with one excep-
tion, has extended from 200 fathoms down to 2507 fath-
oms. The one exception is the Norwegian occurrence at
50 fathoms.
The single specimen from the eastern Pacific appears
to differ from other described species and is designated:
Fissurisepta pacifica CowAN, spec. nov.
Description of Holotype: Shell delicate, white with very
thin straw coloured surface pigmentation, conical, with
anterior slope slightly convex, and posterior slope slightly
concave; foramen somewhat damaged but apparently al-
most circular and situated slightly behind centre. Sculpture
of small pustules widely spaced, nowhere closer to each
other than 4 or 5 times their diameters (Figure 3) ; ar-
ranged generally in horizontal rows around the shell,
about 6 rows in the height of the shell. On the sides, pus-
tules of adjacent rows arranged so that some diagonal
order is apparent, sloping upward toward the anterior
end. Interior of shell glossy white, septum thin, straight,
transverse, extending obliquely downward at an angle of
20° from the anterior slope for about half the height of
the shell; free margin semilunate. Dimensions of shell:
length 5.1mm; width 3.85 mm; height 2.6mm. Subse-
quent to the preparation of this description the specimen
was broken in transit. The fragments remaining reveal
details of surface sculpture but not overall shape, the fora-
men nor the septum.
Vol. 12; No. 1
The soft parts of the genus have not previously been
described.
The soft parts of the posterior segment of the body of
our specimen are badly damaged but the portions avail-
able for study include the head and the mantle cavity
anterior to the heart. Figure 1 illustrates the general
features of the head. From the ventral aspect the oral
disk is wider than long. There are two pairs of cephalic
tentacles, and in the preserved state the anterior pair is
thick, blunt and turned down alongside the oral disk. The
posterior pair is long and slender. There are no eyes.
The most remarkable feature of the superficial anato-
my is that of the ctenidia. Cox (1964, p. 194) states that
all Recent species belonging to the suborder Pleurotomari-
ina, with the single exception of the right ctenidium of
Scissurellidae, have bipectinate ctenidia. Our specimen
Figure 1
Fissurisepta pacifica Cowan, spec. nov.
Ventro-lateral view of head end.
has disproportionately large gills that occupy almost the
entire anterior mantle chamber and both gills are mono-
pectinate (Figure 2). The main axis of each ctenidium is
external and it, as well as most of the length of each
branchial filament, is fused with or adherent to the mantle.
The posterior filaments are free for their terminal thirds.
This is thus the only Pleurotomariine known to have
symmetrically monopectinate ctenidia. Other distinctive
anatomical features should be sought in any fresh speci-
mens that become available.
Type Material: A single known specimen number 6522,
Cowan Collection, University of British Columbia, will be
THE VELIGER
Page 25
Figure 2
Fissurisepta pacifica Cowan, spec. nov.
Dorsal view of anterior third, showing ctenidia as seen through
transparent mantle.
deposited in the type collection of the National Museum
of Canada as N. M. C. Number 45744.
Type Locality: West north west of Triangle Island, Brit-
ish Columbia, 51°09’ N Latitude, 130°09’ W Longitude,
haul number 63-214 of the Fisheries Research Board of
Canada, Cowan station 863; 470 - 480 fathoms. Collected
by D. B. Quayle, September 11, 1964.
Discussion: Fissurisepta pacifica is generally similar in
shell details to EF soyoae Kuropa as described and figured
by him in 1951. It differs, however, in being relatively
flatter, with the height about two thirds of the width
rather than equal to it. It differs also in the number and
arrangement of the tubercles on the shell, which in F
soyoae are shown to be numerous and in well organized
rows of closely placed studs arranged obliquely, whereas
in FE pacifica they are widely separated, few in number
and in generally horizontal rows.
This must be a scarce species as just one specimen has
been taken at about 30 stations below 200 fathoms worked
by us in the past 3 years.
Page 26
An examination of descriptions of other species hither-
to assigned to this subgenus suggests that some detailed re-
examination is necessary. The type species Fissurisepta
papillosa SEGUENZA is known only as a fossil but our
species subscribes to the shell details as does EF soyoae.
Puncturella (Fissurisepta) Wooprinc, 1928 differs in
such essential details of shell structure as thickness, height,
external sculpture and in the size of the shelf. It may well
be improperly assigned to Fissurisepta.
The unique features of the ctenidia of Fissurisepta pa-
cifica lead me to conclude that a separate generic desig-
nation is necessary to recognize the degree of departure
from the characteristics of representatives of Puncturella.
Accordingly, on the assumption that the other species at-
tributed to the subgenus Fissurisepta possess or possessed
the same ctenidial structure, I suggest that Fissurisepta
be accorded generic rank.
Figure 3
Fissurisepta pacifica Cowan, spec. nov.
Surface detail of holotype.
The definition of the genus Fissurisepta can be stated:
Prosobranch, Zeugobranch gastropods of small size with
tall-conical shells in which the aperture is apical. The
septum large, about half the height of the shell and pas-
sing straight from side to side. External sculpture of the
shell variable, consisting of either delicate vertical striae
or of minute, widely separated studs in horizontal or
oblique rows.
Animal with 2 pairs of cephalic tentacles, the posterior
long and slender, no eyes, ctenidia paired, symmetrical
and monopectinate, attached for most of their length to
the inner surface of the mantle.
All but one of the living and fossil forms described
have been taken from the northern oceans. There is, how-
ever, a New Zealand Fissurellid, Puncturella manawata-
whia Powe Lt, that, from published descriptions seems
THE VELIGER
Vol. 12; No. 1
closely similar. It would be interesting to study this species
in more detail.
ACKNOWLEDGMENTS
Dr.James H. McLean, Los Angeles County Museum of
Natural History assisted with the preliminary identifica-
tion and provided the photographic figures; the anatom-
ical figures are the work of Margaret Jensen.
LITERATURE CITED
CrarkE, ArTHUR H., Jr.
1961. Abyssal mollusks from the South Atlantic Ocean.
Bull. Mus. Comp. Zool., Harvard, 125 (12): 345 - 387; 4 plts.
Cox, Lestie REGINALD
1960. | Gastropoda/General characteristics of Gastropoda
In: R.C. Moore, Treatise on invertebrate paleontology Prt. I,
vol. 1: 84 - 169; figs. 51 - 88E
Dati, WiLL1AM HEALEY
1889. Reports on the results of dredging, . . . in the Gulf of
Mexico (1877-78) and in the Caribbean Sea (1879-80), by
the U. S. Coast Survey Steamer “Blake”, . . . Report on the
Mollusca, pt. 2, Gastropoda and Scaphopoda. Bull. Mus.
Comp. Zool. 18 (2) : 1 - 492; plts. 1 - 40
1927. Small shells from dredgings off the southeast coast of
the United States by the United States Fisheries Steamer
“Albatross” in 1885 and 1886. Proc. U. S. Nat. Mus. 70:
1- 134
DAUTZENBERG, PHILIPPE & HENRI FISCHER
1927. Résultats comparat. scientif: accomplies sur son yacht par
Albert 1** de Monaco. 77: 224; plt. 7, fig. 16
FaRFANTE, ISABEL PEREZ
1947. The genera Zeidora, Nesta, Emarginula, Rimula, and
Puncturella in the Western Atlantic. Johnsonia 2 (24):
93 - 148
JEFFREYS, JOHN Gwyn
1882. On Mollusca procured during the “Lightning” and
“Porcupine” expeditions, 1867-70 (Part V). Proc. Zool.
Soc. London for 1882: 656 - 687; 2 plts.
Kuropa, ToKuUBEI
1951. _— Illustrated catalogue of Japanese shells. No. 17: Fissur-
ellidae, p. 116
OKUTANI, TAKASHI
1964. | Report on the archibenthal and abyssal gastropod Mol-
lusca mainly collected from Sagami Bay and adjacent waters by
the R. V. Soyo-Maru during the years 1955 - 1963. Journ.
Fac. Sci., Univ. Tokyo, Sec. II, 15 (3): 371 - 447
PowELL, ARTHUR WILLIAM BADEN
1937. | New species of marine mollusca from New Zealand.
Discovery Reprts. 15: 153 - 222; 11 plts.
SEGUENZzA, G.
1863. Paleontologia malacologia dei terreni Terziarii del dis-
tretto dei Messina ... Fanuglia Fissurellidi. Amali Accad.
Aspir. Naturalisti, Naples, ser. 3, 2: 83 - 86
Wooprinc, WENDELL PHILLIPS
1928. Miocene mollusks from Bowden, Jamaica. Part 2: Gas-
tropods and discussion of results. Carnegie Inst. Washington,
Publ. 385: i- vii + 1-564; 3 text figs.; plts. 1-40 (28 Nov.)
peng
Vol. 12; No. 1
THE VELIGER
Page 27
Seasonal Gonadal Changes of Adult Oviparous Oysters
in Tomales Bay, California
BY
CARL J. BERG, Jr.
Pacific Marine Station, Dillon Beach, California ':?
(Plates 1 to 3; 1 Text figure)
INTRODUCTION
Two SPECIES OF oviparous oysters, Crassostrea virginica
(GmetIn, 1791), the Eastern oyster, and Crassostrea
gigas (THUNBERG, 1793), the Japanese oyster, were intro-
duced into Tomales Bay, California for commercial pur-
poses. Because these oysters do not reproduce in its waters,
shipments of adult oysters or oyster spat must be period-
ically transplanted into this bay. It is the purpose of
this study to compare the seasonal gonadal changes which
occur in the two species of adult oysters in Tomales Bay
with one another, and to compare them with similar
changes occurring in the same species of oysters in other
regions. The differences in gonadal changes will be related
to variations in environmental conditions and to specific or
racial characteristics.
Tomales Bay is located on the Northern Californian
Coast, approximately 40 miles north of San Francisco.
The oysters used in this study were kept at Tomales Bay
Oyster Company, which is situated in the cove between
Millerton and North Double Point, near the headwaters
of the bay. This area has long been the site of oyster
culture. The native or Olympia oyster (Ostrea lurida
CarPENTER, 1864) was the first oyster species grown in
Tomales Bay to be shipped to the markets of San Fran-
cisco. However, upon completion of the transcontinental
Central Pacific Railroad in May 1869, it was possible for
the first time to ship live adult Crassostrea virginica and
its seed to the Pacific coast. In 1875, 17 carloads of C.
virginica were laid out near Millerton Station in Tomales
Bay (TownsEnp, 1893). Cultivation of C. virginica in
Tomales Bay has continued intermittently since that time.
‘ Submitted in partial fulfilment of the requirements for the
Degree of Master of Science in Marine Science at the University
of the Pacific, Stockton, California.
2 Present address: Department of Zoology, University of Hawaii,
Honolulu, Hawaii 96822
The tidelands of Tomales Bay Ovster Company were also
the first areas in California to be used for the culture of
Japanese oysters (C. gigas), which were introduced into
Puget Sound in 1902, and later, in 1928, into Tomales Bay
(BarreTT, 1963). Both C. virginica and C. gigas have
failed to reproduce in Tomales Bay and continued impor-
tation of adult oysters or seed has been necessary.
Because of their commercial value, there have been
numerous attempts at the artificial introduction of oysters
into areas lacking natural populations, or in which the
native oyster is of little economic importance. The oysters
of the genus Crassostrea Sacco, 1897, are more often
used as introduced species than are those of the genus
Ostrea LINNAEUS, 1758, since they have a greater ability
to survive in extreme and varying environments. Cyrass-
ostrea virginica has been introduced into the oyster beds
of Britain, the Pacific Coast of North America, and the
island of Oahu, Hawaii. Crassostrea gigas has likewise
been introduced into the oyster beds of the Pacific Coast
of North America; Oahu, Hawaii; also into Melbourne
Harbour, Australia; Mobile Bay, Alabama; and Barnstable
Bay, Massachusetts. The Portuguese oyster, C’. angulata
(Lamarck, 1819) has taken over all of the French oyster
beds and now also lives, but does not reproduce, in
British beds. One noteworthy and successful introduction
of oysters of the genus Ostrea took place in the waters of
Boothbay Harbor, Maine, where LoosANorF (1955) had
transplanted European oysters (Ostrea edulis LINNAEUS,
1758) in 1949.
In many of the attempts to introduce a specics of
oyster to a new location, the animals may have survived
and grown well, but failed to reproduce. To determine
the reason for the failure of the species to reproduce,
one must be familiar with the seasonal gonadal changes
which occur in the oysters in their native environment.
There have been studies on gonadal changes in each of
the commercially valuable species of oysters. Crassostrea
Page 28
virginica is the most extensively studied species of the
oviparous oysters (Hopkins, 1931; Loosanorr, 1932,
1942, 1965; LoosaNorF & ENGLE, 1942; BuTLER, 1949;
KENNEDY & BATTLE, 1964; SEVILLA & Monpracon, 1965).
Crassostrea gigas has had little work done on its gametic
activity (Imai et al., 1950; Imar & Sax1, 1961) and the
same is true with the other oviparous oysters: C. commer-
cialis (IREDALE & RouGHLEy, 1933) (RouGHLEy, 1933;
Cieanp, 1947); C. angulata (BARcETON, 1942, 1943) ;
and C. madrasensis (PRESTON) (ANOoNYMouS, 1950).
Studies have also been made of gonadal changes in the
larviparous oysters: Ostrea edulis (Orton, 1927, 1931,
1933; Core, 1942) and O. lurida (Cor, 1931, 1932).
Even less work has been done on the seasonal gonadal
changes in oysters which have been introduced into new
areas. LoosanoFF (1962a) gave a detailed description of
the changes which occurred in the gonads of O. edulis
introduced into Boothbay Harbor, and briefly mentioned
the gonadal changes which occurred in C. gigas trans-
planted into Milford Harbor (LoosanorF & Davis, 1963).
Ga.tsorF (1929) and KaTKaNSky & SPARKS (1966)
discussed the gonadal changes and sex ratios which oc-
curred in C. gigas cultured in the waters of the State of
Washington, and Evsey (1932, 1933, 1934) described the
changes in C. gigas in the waters of British Columbia.
Ostrea lurida is the only other species in which the sea-
sonal gonadal changes have been thoroughly studied in
both the native environment (Cog, 1931, 1932) and in the
area to which it was introduced (Hort, 1933).
The introduction of oysters into a new environment is
valuable for both scientific and commercial reasons. Al-
though it has long been known that Crassostrea virginica
and C’.. gigas do not reproduce in Tomales Bay, no one
has ever done a detailed study of the reproductive cycle
or the seasonal histological changes in the gonads. This
study, therefore, extends the knowledge concerning an
oyster’s adaptation to its new environment and may help
to define the reasons for the failure of these two species
of oysters to reproduce in Tomales Bay.
MATERIALS ann METHODS
The oysters used in this study were obtained from Long
Island Sound and from Canadian waters through the
courtesy of the Department of Fish and Game of the
State of California, Dr. Victor L. Loosanoff and Dr. Ed-
mund H. Smith of the Pacific Marine Station. Four hund-
red and fifty 2-year old Crassostrea gigas of Canadian ori-
gin were taken from the mud-flats at Tomales Bay Oyster
Company where they had been raised. They were then
placed in wire-mesh trays and suspended from racks at
THE VELIGER
Vol. 12; No. 1
the Oyster Company on October 20, 1966. A similar
number of 2-year old C. virginica was received on Octo-
ber 26, 1966, and was placed in identical adjacent trays.
Fifteen oysters of each species were collected at bi-weekly
intervals for 15 months. An additional sample of 13 C.
virginica was obtained on October 5, 1966, from a stock
which had been previously kept at Tomales Bay Oyster
Company.
The height (distance between the umbo and the ventral
valve margin) and length (distance between anterior and
posterior valve margins) of each oyster were measured
with calipers before the oyster was opened. A tissue sample
was taken from the gonadal area near the labial palps in
conformity with the practice of the biological labora-
tories of the United States Bureau of Commercial Fish-
eries. This gonadal tissue was fixed in Bouin’s solution,
infiltrated with 52.5° C paraffin, sectioned at 10 B, and
stained with Heidenhain’s iron hematoxylin and aqueous
eosin Y, using standard procedures. The tissues were then
microscopically analysed for sex and state of gametogen-
esis.
Hydrographic data were collected on each sampling
date; they included measurements of salinity, hydrogen-
ion concentration, and turbidity of water samples taken
one foot above the bottom by means of a Frautschy-bottle
sampling device. Sub-surface water temperatures were
taken by suspending a thermometer over the bottom.
Surface-water hydrographic data were obtained from rec-
ords kept at the Pacific Marine Station.
Large bags of sun-bleached oyster shells were suspended
near the trays of oysters for 2-week periods from May 4,
1967 until October 19, 1967. Fifty shells were chosen at
random from each bag and examined for oyster spat.
During the same period plankton tows were taken over
the beds of Tomales Bay Oyster Company and examined
for bivalve larvae. All oyster larvae were identified by
length-width measurements after the method of Loosan-
oFF, Davis & CHANLEY, 1966.
OBSERVATIONS
A detailed description of the sequence of events in the
development of functional gametes has been given by
LoosanoFF (1942) for Crassostrea virginica. Since the
seasonal gonadal changes observed in both C. virginica
and C. gigas in Tomales Bay differed little from those
described, except with regard to timing (LoOOSANOFF &
EncLE, 1940; Loosanorr, 1965), only a résumé of the
gonadal changes will be given here. It is most convenient
to describe the seasonal gonadal changes which occurred
Vol. 12; No. 1 THE VELIGER Page 29
Salinity %,
Temperature in ° C
Co)
ONDJFMAMJJAS ON D
Month
Figure 1
Water temperature and salinity at Tomales Bay Oyster Company
from October 1966 to December 1967
in the two species separately and in the order in which
they were observed.
Specimens of Crassostrea virginica were obtained from
Tomales Bay Oyster Company on October 5, 1966, where
they had been kept for at least a year. The small gonadal
follicles of these oysters contained only indifferent sex cells
and were scattered in the large masses of connective tissue.
This “indifferent stage” (LoosanorrF, 1942, p. 203) was
characteristic in the months of September, October, and
November of 1967.
Page 30
The remaining samples of gonadal tissue were collected
from Crassostrea virginica transplanted into Tomales Bay
Oyster Company beds on October 26, 1966. Slight ga-
metic activity ended the indifferent stage of some of the
oysters during the months of November and December.
Primary and secondary gametogonia started to develop
along the follicular walls, making sex determination pos-
sible. The follicles began to expand gradually.
A marked increase in the size of the follicles and in
the acceleration of maturation of the gametes was no-
ticed in late December and January. Ovocytes began
to fill the follicles and spermatids were already devel-
oping. The follicles showed rapid growth and ramification.
Continued maturation of the gametes took place in
February and March, concurrently with the disappearance
of the voluminous vesicular connective tissue. In some
instances, mature ova and spermatozoa were present.
By the sixth of April, most of the gonads were packed
with ripe gametes, some of which nearly filled the ciliated
genital ducts (Plate 1, Figures 1 and 2). However, a
few cells at the earlier stages of gametogenesis were pres-
ent on the follicular walls. This state of maximum ripe-
ness was maintained throughout the spring. Although a
few oysters were partially spawned beginning at the
middle of April, mass spawning did not occur until the ©
middle of June and continued until the middle of July.
On July 13, 87% of the sample was completely spawned.
This coincided with the time of greatest rate of increase
in water temperature (Text figure 1).
The gonads of spawned oysters were characterized by
the absence of mature gametes and the shrunken appear-
ance of the follicles (Plate 1, Figures 3 and 4). Great
numbers of phagocytic cells were present, both inside the
lumina of the follicles and around the outside walls. All
unshed gametes are devoured by these cells. The lumina
of the follicles were being closed by elongation and the
shrinking of follicular tissue. Simultaneously, the cells of
THE VELIGER
Vol. 12; No. 1
the vesicular connective tissue proliferated, filling all inter-
follicular spaces. Resorption of gametes continued from
the post-spawning stages in July until the first week of
November, and a few follicles were observed to contain
mature gametes being phagocytized as late as December.
However, most gonads were in the indifferent stage by
September.
All samples of gonadal tissue from Crassostrea gigas
were collected from oysters which had been raised in
Tomales Bay from the imported seed. Although the sea-
sonal gonadal changes which were observed in C. gigas
resembled those of C. virginica, there were differences
between species in timing and in the homogeneity of the
sample. The indifferent stage of gametogenesis was char-
acteristic of the months of November and December.
The little gametic development and proliferation of
the follicles which occurred in December and January is
the only activity which might be characterised as the “sex-
differentiation stage” (Loosanorr, 1942, p. 203).
Very rapid maturation of the gametes took place in
January and February, with a great expansion of the
follicles. March was the month of greatest maturation and
proliferation. By the eleventh of March, ripe ova and
spermatozoa were present in a few of the follicles.
From April 6 until July 27, 95% of all the gonads
sampled were filled with ripe gametes (Plate 2, Figures 5
and 6). They retained gametes throughout the spring,
however, because no spawning took place until after the
twenty-seventh of July. Between the dates of July 27 and
August 10, mass spawning of Crassostrea gigas occurred.
This coincided with the warmest water temperatures to
that date (Text figure 1). Again, the follicles of the
spawned gonads were shrunken and devoid of ripe gam-
etes (Plate 2, Figures 7 and 8).
Resorption and cytolysis of the unshed gametes started
immediately after spawning was completed and con-
Explanation of Plate 1
Figure 1: Section of gonad of ripe female Crassostrea virginica
collected April 6, 1967 (X 125)
Figure 2: Section of gonad of ripe male Crassostrea virginica
collected April 6, 1967 (X 125)
Figure 3: Section of gonad of spawned female Crassostrea virginica
collected July 13, 1967 (X 125)
Figure 4: Section of gonad of spawned male Crassostrea virginica
collected July 13, 1967 (X 125)
Explanation of Plate 2
Figure 5: Section of gonad of ripe female Crassostrea gigas
collected April 6, 1967 (X 125)
Figure 6: Section of gonad of ripe male Crassostrea gigas
collected April 6, 1967 (X 125)
Figure 7: Section of gonad of spawned female Crassostrea gigas
with a few ova remaining in gonad tubules.
collected August 10, 1967 (X 125)
Figure 8: Section of gonad of spawned male Crassostrea gigas
showing spermatozoa remaining in gonad tubules.
collected August 10, 1967 (X 125)
[BERG] Plate 1
Tue VELIGER, Vol. 12, No. 1
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Vol. 12; No. 1
tinued until December. Many spermatozoa and ova were
found being phagocytized, some until late December.
In addition to observing the gonadal changes, the sex
ratio was also noted. Frequently, however, the sex could
not be determined since the gametes were not yet differ-
entiated. In a sample of 387 Crassostrea virginica, 139
were male, 114 were female, and 132 were undifferenti-
ated. Two morphological hermaphrodites were found.
Both contained follicles packed with spermatozoa and a
few cytolized ova (Plate 3, Figure 9). In a sample of 382
C. gigas 100 were male, 194 were female, and 87 were
undifferentiated. Only one hermaphrodite was found for
this species. Unlike those observed in the sample of C.
virginica, the follicles of the hermaphroditic C. gigas con-
tained both male and female gametes in various stages of
development (Plate 3, Figure 10).
The hydrographic conditions observed in the bay are
presented in graphic form (Text figure 1). The range of
hydrogen-ion concentration was not great enough (7.70
to 8.16) to affect gonadal changes, neither did it exhibit
seasonal variances. Hence, these data are not presented in
this article. The turbidity determinations are not presented
either, although the turbidity of the water showed distinct
seasonal variations. Both temperature and salinity may
affect gametogenesis (KINNE, 1963, 1964; LoosaNnorr,
1945, 1948). Text figure 1 presents temperature and sa-
linity data in graphic form so that seasonal irends may be
more apparent.
The difference between surface and bottom salinity
samples is significant and noteworthy since the oysters
were exposed to surface water at low tides. The extreme
range in salinities is due to the effect of fresh-water runoff
during the winter and spring, and to evaporation during
the summer. Salinities in excess of 32%, prevailed from
July to December. Throughout the winter and spring,
however, the salinity remained around 25%.
The water temperature at the oyster beds exhibited
similar seasonal fluctuations. A trend of rising tempera-
tures starts in January after a brief cold period and con-
tinues until June. The water temperature remains close to
20°C during June, July, August and September. The
water then begins to cool to its lowest temperature at
the end of the year. Rarely did the temperature remain
below 10°C for more than a few days, neither did it
range much above 20° C. All hydrographic observations,
including temperature determinations, were made at
approximately mid-tide. More extreme temperatures may
exist during slack tides, but thermograph records for that
area showed that the water temperatures remain markedly
constant throughout the tidal cycle.
Finally, few Crassostrea larvae were collected by the
plankton tows and few spat were found on shells. Some
THE VELIGER
Page 31
straight-hinge stage larvae were found in the plankton
samples of August 24 which had general appearances and
dimensions similar to those reported for C. virginica and
C. gigas (LoosanorF, Davis & CHANLEY, 1966). Although
bivalve larvae were collected in every sample, no others
approximated the proper size or shape.
A total of 6 oyster spat was found set on shells; one on
July 13, and 5 on August 10. Three of those found on
August 10 had settled on the inside of the right valve of
an oyster which was gaping. The inside of the shell was
free of mud, although the outside was covered. Through-
out the summer, all shells in the wire-mesh bags were
found to be covered with mud after 2 weeks in the water.
It is surprising that any larvae were able to set on such
shells.
DISCUSSION
Gametogenesis and spawning of oysters are directly cor-
related to water temperatures. LoosanorF & Davis in
1952 have shown the temperature and time requirements
to condition Crassostrea virginica to spawn. Their experi-
ments showed that 10° C was not high enough to induce
gametic activitics. However, they report ripening and
spawning for oysters when the temperature had reached
only 15°C. Later experiments (Loosanorr, 1958, 1969)
proved that maturation of gametes was possible after 68
days of conditioning at 12° C. The water temperatures at
Tomales Bay Oyster Company remained above 12°C
throughout the year, except for short periods of time in
December and January. During the summer, the water
temperature was not as high as those reported for either
Long Island Sound or the Miyagi Prefecture, Japan, from
where the oysters originally came. The differences in
seasonal temperature fluctuations between Tomales Bay
and the native environments of the oysters were respon-
sible for the variances in gonadal changes (Table 1).
Low salinities which may have an influence on gonad
development (LoosanorFr, 1948; 1952), were not en-
countered for extended periods of time, thus they did not
seem to affect gonadal changes in oysters in Tomales Bay.
A comparison of seasonal gonadal changes between
Crassostrea virginica and C. gigas must take into account
differences between the species and racial variations
within each species. As mentioned previously, the seasonal
gonadal changes observed in both C. virginica and C.
gigas differed little from each other, except for timing.
The differences in timing are clearly portrayed in Table 1.
The only time of agreement in stages of gametogenesis
was from the first week in April, when approximately 757%
of the samples from both species contained ripe ova and
Page 32
THE VELIGER
Vol. 12; No. 1
Table 1
Periods of Basic Gonadal Stages in Crassostrea virginica and Crassostrea gigas in Different Geographic Areas.
Crassostrea virginica
Stage Tomales Bay Long Island
California Sound
Spring January
Development and and
February May
Crassostrea gigas
Tomales Bay State of Miyagi Prefecture,
California Washington Japan
February May
and —
March June
First April June April May May
Ripening ‘
Spawning through
August
From Post- From Post-
Resorption Spawning to Spawning to
October October
Undiffer- September August
entiated through and
November September
Fall November October
Gametogenesis and and
December November
December
Inactive None to
April
sperm, until the end of June when most of C. virginica
had spawned. During this period, no C. gigas spawned,
but a few of C’. virginica had partially spawned and some
were undergoing resorption. Mass spawning of C. virgini-
ca was completed at least two weeks before C. gigas
spawned, and it extended over a greater period of time
than did the 2-week mass spawning of C. gigas. Since
the shed gametes of both species can induce spawning in
the males of either species (GaLTSoFF, 1964), it is sur-
prising that C. gigas did not spawn simultaneously with C.
virginica. A combination of thermal and chemical re-
quirements is often needed to induce spawning. These
requirements were obviously not met for C. gigas until a
later date.
Not all of the oysters of either species had spawned
completely. More of Crassostrea gigas contained ripe ova
and spermatozoa, and retained them until later dates,
than did C. virginica. Resorption of unshed gametes con-
tinued into December for C’. gigas. During and following
resorption, the gonadal follicles were in the undifferenti-
ated stage. This stage was halted with the advent of
gametogenesis. In C. virginica, gametogenesis started in
November, one month earlier than in C. gigas. Matura-
tion proceeded gradually until after the first of the year
for both species. In late December, January, and February,
July July August
and and and
August September
From Post-
Spawning to —
December
From Post-
Spawning to
December
October
through
November
and
December
December
February
through
April
the gametes of C. virginica showed a great acceleration
in maturation. This acceleration, termed “spring develop-
ment” by LoosanorF (1942, p. 198), again occurred one
month earlier in C. virginica than in C. gigas. However,
C. gigas showed a greater burst of gametic activity in
March. Thus, both species reached sexual maturity at the
same time. In general, C. gigas has a higher temperature
requirement for the initiation of both gametogenesis and
spawning. This is evident by the slower development and
later spawning dates. Moreover, unshed gametes are re-
tained by C. gigas for longer periods of time before being
resorbed. Similar differences were observed between C.
virginica and C. gigas transplanted into Milford Harbor
(LoosanorF & Davis, 1963).
A series of observations on seasonal gonadal changes of
two other bivalve species was conducted concurrently
with this study (LEonarD, 1969). Ostrea edulis, trans-
planted into Tomales Bay Oyster Company beds, and
Pododesmus cepio (Gray, 1850), the native rock jingle,
both showed gonadal changes at dates which correspond
well with those observed for Crassostrea virginica and C.
gigas and with the seasonal fluctuations in temperature.
A comparison of the timing of the gonadal changes of
Crassostrea virginica in Tomales Bay with those reported
for it in Long Island Sound (Loosanorr, 1942; 1965)
Vol. 12; No. 1
emphasizes tiie effect of temperature upon gametogen-
esis and spawning (Table 1). Spawning occurred in both
locations at the end of June when the water temperature
was about 20° C. Post-spawning resorption continues into
October in both locations, but the undifferentiated stage
is being completed in Long Island Sound in September,
and not until November in Tomales Bay.
Fall gametogenesis extends from October to December
in Long Island Sound but occurs one month later in
Tomales Bay. Whereas Long Island Sound oysters exhibit
no gametic activity from December until April or May,
oysters in Tomales Bay proceeded to carry out gameto-
genesis through the winter, with the period of fastest ma-
turation occurring in February. In April, when oysters in
Tomales Bay were already ripe, Long Island Sound oysters
were developing only primary and secondary gametocytes.
During the entire spring, oysters in Tomales Bay contain
ripe gametes, while the gametes in Long Island Sound
oysters are just maturing. In both areas, the gametes are
ripe in June.
To reiterate, the major difference in gonadal changes
between Crassostrea virginica in Long Island Sound and
those transplanted into Tomales Bay is that the follicles
of the latter do not exhibit an inactive stage; rather they
continue to develop from November until they are mature
in April. The ripe gametes then remain in the follicles
until spawning occurs. This is probably due to differences
in winter temperatures of the two areas.
There is little information on the seasonal gonadal
changes of Crassostrea gigas. Although a few authors
mention the temperature at which spawning occurred,
none describe the histological changes taking place in the
gonads prior to spawning. Spawning is recorded for
temperatures below 20°C (Fuyrra, 1929; Hopxins,
1936; ScHarFEeR, 1938; CuHapMaN & EsvetpT, 1943),
which is well within the range encountered in Tomales
Bay. Imar & Saki (1961) report that spawning takes place
in Miyagi strains of C. gigas late in August and early in
September, although the gonads were ripe as early as
May. Amemiya (1928), however, reports that spawning
of northern province Japanese oysters may take place in
May and June. The timing of gametogenesis and spawning
of C. gigas in Tomales Bay best corresponds with the data
from Imari & SAKI (op. cit.) but seems to be one month
ahead at all stages.
Crassostrea gigas raised in the State of Washington
also exhibit gonadal changes and spawning at tempera-
tures almost identical with those observed in Tomales Bay.
Spawning is reported to occur during the last part of
July and the first part of August when the water tempera-
tures ranged between 18° C and 20°C (Scuaerer, 1938;
CHAPMAN & EsveLpT, 1943). This coincides perfectly
THE VELIGER
Page 33
with the observations of spawning in Tomales Bay. In
the follicles of C. gigas in Washington waters, resorption
of the unshed gametes continues through December
(Ga.tsorF, 1929). The follicles begin to proliferate in
December and January (KaTansky & Sparks, 1966),
but then gametic activity appears to be arrested. Re-
sumption of maturation of the gametes takes place in
April or May, with ripe ova and active sperm again
present by May (GaLTsorr, op. cit.; CHAPMAN & Es-
VELDT, op. cit.). The inactive wintering stage is similar
to that of C. virginica in Long Island Sound. As mentioned
previously, C. gigas in Tomales Bay does not become in-
active; rather the gametes mature most during the months
of January, February, and March.
It is apparent that Crassostrea gigas in Tomales Bay
meets with seasonal temperature fluctuations which are
similar to those in the State of Washington, where they
were first introduced. Normal gametogenesis and spawn-
ing occurs in all three areas. Only in Japan, however, is
the combination of all environmental factors consistently
suitable for the setting of oyster larvae. Some setting of
commercial value also occurs occasionally in Washington
and Canadian waters. In fact, the oysters used in this
study were from Canadian seed. There have been no
reports of a large set of either C. gigas or C. virginica
in Tomales Bay, although one naturally set C. gigas was
found. Factors other than temperature, salinity, and hy-
drogen-ion concentration are probably involved, since
these factors correspond well with those found in the
oyster’s native environment. Excessive turbidities were
noticed throughout the summer. These probably had a
deleterious effect upon both the larvae and the places
available for setting. A more complete discussion of the
effect of the environment upon the larvae will be pre-
sented at a later date.
LoosaNnoFF (1962b) has shown the effect of turbidity
upon oyster larvae, and LoosANoFF & ToMMERS (1948)
have suggested that there exist races of C. virginica
which exhibit different abilities to survive high turbidi-
ties. Furthermore, races based upon the oysters’ adapt-
abilities to an environment, shell characters, and temper-
ature requirements for breeding have been described for
both C. virginica and C. gigas (STauBER, 1950; LoosANOFF
& NoMEjKo, 1951; Imar & Saxt, 1961; Loosanorr, 1969).
Oysters from northern latitudes have been shown by these
authors to have a much lower temperature requirement
for the development of gametes and the induction of
spawning. The results of this study showed great differ-
ences within the population of C. virginica dredged from
Long Island Sound and transplanted into Tomales Bay.
In any one sample, the oyster gonads would show a
continuum through two or three stages of development.
Page 34
Some oysters ripened and spawned much earlier than
most; others much later. In addition, resorption of unshed
gametes proceeded at greatly varying rates. Some oysters
contained ripe ova and spermatozoa even in December.
The population of C. gigas did not show as great variance,
probably because the oysters were collected as seed from
a small area and then transplanted into Tomales Bay. The
greater homogeneity of the population indicates that
only one race was observed. Some individual differences
were noted, but this should be expected. ‘The oysters used
in this study came from the northern limits of the distri-
bution of each species. If more southern oysters were
used, the environmental conditions in Tomales Bay would
not have favored spawning. In all attempts at intro-
ducing a species into a new area, consideration must
be given to the racial characteristics of the oysters.
Animals with the lowest temperature requirements are
best suited for such experiments.
Recent findings support the earlier observations that
crowded conditions of oysters might favor the develop-
ment of one sex or the other. In crowded conditions,
Crassostrea virginica are reported to show a population
trend toward the male phase (BuRKENROAD, 1931; NEED-
LER, 1934) and C’. gigas to the female phase (KATANSKY
& SPARKS, 1966). However, experimental evidence (V. L.
LoosanorF, MS) has shown that crowded conditions
did not affect sex determination in adult C. virginica. The
samples taken for this study indicate that a female phase
may be favored by C. gigas in crowded conditions, but
that C. virginica showed approximately equal sex ratios.
The number of hermaphrodites found is also within the
normal range. (Gattsorr, 1964). In Crassostrea gigas
0.26% were hermaphrodites (1 out of 382) and in C.
virginica 0.52% (2 out of 387). The few ova present in
the ripe male follicles of C. virginica were cytolized
(Plate 3, Figure 9), but both the ova and the spermatozoa
in C. gigas appeared normal (Plate 3, Figure 10), with
primary and secondary spermatocytes present. Therefore,
the hermaphrodites observed in C.. virginica would best be
called morphological, but the one C. gigas was probably a
functional hermaphrodite. All of the hermaphrodites
were collected in the spring during the period of greatest
gametic activity. It is unlikely, therefore, that the gametes
had been carried over from the previous season.
A few major conclusions have been drawn from this
study of the seasonal gonadal changes in Crassostrea
THE VELIGER
Vol. 12; No. 1
virginica and C’. gigas introduced into Tomales Bay, Cali-
fornia. Differences in the gonadal changes between in-
dividuals of each species, between two species, and
between the oysters in Tomales Bay and those of the same
species in their native environment are attributed to
individual, racial, specific, and environmental variations.
The sample of C. gigas was racially more homogeneous
than that of C. virginica, and it required warmer temper-
atures to initiate gametogenesis and spawning. During the
annual cycle, one continuous period of gametic activity
took place in both species. There was no inactive stage as
is usually found in most species of Crassostrea in their
native environments, because the water temperatures at
Tomales Bay Oyster Company were favorable enough
throughout the year to encourage gametogenesis and
growth. The inability of the two species to propagate in
Tomales Bay is not due to a failure to spawn, but rather
to a failure of the larvae to reach the settled spat stage.
ACKNOWLEDGMENTS
I wish to thank Dr. Victor L. Loosanoff for the invaluable
guidance and encouragement he gave me throughout the
course of this study. My thanks are also extended to Dr.
Edmund H. Smith, Dr. John S. Tucker, and Dr. Walter
Narchi for their helpful advice and assistance.
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Barrett, ELINoRE M. >
1963. The California oyster industry. Calif. Dept. Fish
& Game, Fish Bull. 123: 103 pp.; 32 text figs.
Explanation of Plate 3
Figure g: Section of gonad of a hermaphroditic Crassostrea vir-
ginica containing both spermatozoa and cytolized ova.
collected May 18, 1967 (X 125)
Figure 10: Section of gonad of a hermaphroditic Crassostrea gigas
containing both spermatozoa and ova
collected April 6, 1967 (X 125)
TuHeE VELIcER, Vol. 12, No. 1 [Bers] Plate 3
Figure 10
i a we = _
i" ae 7,
UP an
: 7 i
= = , <
> 4+ - ‘
t
a
¥ *
_
i
= = = i
= f
2,
/
J
:
~
7
:
Vol. 12; No. 1
THE VELIGER
Page 35
BuRKENROAD, Martin D.
1931. Sex in the Louisiana oyster, Ostrea virginica. Sci-
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1943. The spawning and setting of the Pacific oyster (Ostrea
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1947. | Some observations on the cytology and oogenesis in the
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1932. Japanese oysters breed in Ladysmith Harbour, B. C.
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1933. | Oysters in British Columbia. Biol. Brd. Canada Bull.
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1931. Factors influencing the spawning and setting of oysters
in Galveston Bay, Texas. Bull. U.S. Bur. Fish. 47 (3):
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1933. On the development of the Olympia oyster, Ostrea luri-
da CarPENTER, transplanted from United States to Japan.
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Imatr, TAKEO, Masayosui Hatanaka, RyunHeEtr Sato, Secu SAKI,
& Ryoco YuKI
1950. Artificial breeding of oysters in tanks.
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1961. Study of breeding of Japanese oyster, Crassostrea gigas.
Tohoku Journ. Agr. Res. 12 (2): 125-171; plts. 1-2; 16 figs.
24 tables
Tohoku
KATKANSKY, STANLEY C. & ALBERT K. SPARKS
1966. Seasonal sexual patterns in the Pacific oyster Crassostrea
gigas in Washington State. Fish. Res. Pap., Wash. Dept.
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KENNEDY, AusTINA & HELEN I. BATTLE
1964. Cyclic changes in the gonad of the American oyster C.
virginica (GMELIN). Canad. Journ. Zool. 42 (2) : 305 - 321;
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KINNE, OTTo
1963. The effects of temperature and salinity on marine and
brackish water animals. I. Temperature. Jn: Oceanography and
Marine Biology. H. Barnes (ed.). vol. 1. George Allen « Unwin
Ltd., London; pp. 301 - 340; 7 text figs.; 3 tables
1964. The effects of temperature and salinity on marine and
brackish water animals. II. Salinity and temperature-salinity
reactions. In: Oceanography and Marine Biology, H. Barnes
(ed.), vol. 2. Hafner, New York; pp. 281 - 340; 17 text figs.;
14 tables
LEonarD, VERNON KENNETH, Jr.
1969. Seasonal gonadal changes in two bivalve mollusks in
Tomales Bay, California. The Veliger 11 (4): 382-390;
plts. 58 - 60; 4 text figs.; 1 map (1 April 1969)
Loosanorr, Victor Lyon
1932. | Observations on propagation of oysters in James and
Corrotomon Rivers and the seaside of Virginia. Publ. Va.
Comm. Fish. 46 pp.; 17 text figs.
1942. Seasonal gonadal changes in the adult oysters, Ostrea
virginica, of Long Island Sound. Biol. Bull. 82 (2): 195
to 206; plts. 1-3; 12 text figs.
1945. Precocious gonad development in oysters induced in
midwinter by high temperature. Science 102 (2640) :
124 - 125
1948. | Gonad development and spawning of oysters (O. vir-
ginica) in low salinities. Anat. Rec. 101 (4) : 55
1952. Behavior of oysters in water of low salinities. Proc.
Natl. Shellfish Assoc. 43: 135-151; 7 tables
1955. The European oyster in American waters. Sci.
121 (3135): 119-121
1958. Challenging problems in shellfish biology. In: Per-
spectives in marine biology, Part ITV (1956), A. A. Buzzattt-
Traverso (Ed.), Univ. Calif: Press, Berkeley & Los Angeles, pp.
483 - 495
1962a. Gametogenesis and spawning of the European oyster,
Ostrea edulis, in waters of Maine. Biol. Bull. 122 (1):
86 - 94; 15 text figs.
1962b. Effects of turbidity on some larval and adult bivalves.
Proc. Gulf and Carib. Fish. Inst. 14t Ann. Sess., 80 - 95; 2 tab.
9 text figs.
1965. | Gonad development and discharge of spawn in oysters
of Long Island Sound. Biol. Bull. 129 (3): 546-561; 6
text figs.; 3 tables
1969. Maturation of gonads of oysters, Crassostrea virginica,
of different geographical areas subjected to relatively low tem-
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(1 January 1969)
LoosanoFF, Victor Lyon « Harry Cari Davis
1952. Temperature requirements for maturation of gonads of
northern oysters. Biol. Bull. 103 (1): 80-96; 14 text figs.
5 tables
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Loosanorr, Victor Lyon « Harry Cart Davis, continued
1963. Rearing of bivalve mollusks. Jn Advances in marine
biology, vol. 1, FS. Russell (ed.). Academic Press, London,
pp. 1-136; 43 text figs.
Loosanorr, Victor Lyon, Harry Cart Davis & Paut E. CHANLEY
1966. Dimensions and shapes of larvae of some marine mollusks.
Malacologia 4 (2): 351 - 435; 61 text figs.
Loosanorr, Victor Lyon & JAMEs B. ENGLE
1940. Spawning and setting of oysters in Long Island Sound
in 1937, and discussion of the method for predicting the in-
tensity and time of oyster setting. Bull. U.S. Bur. Fish.
49: 217 - 255; 11 text figs.; 18 tables
1942a. Accumulation and discharge of spawn by oysters living
at different depths. Biol. Bull. 82 (3) : 413 - 422; 4 text figs.
Loosanorr, Victor Lyon & CHartes A. NoMEJKo
1951. Existence of physiologically-different races of oysters,
Crassostrea virginica. Biol. Bull. 101 (2): 151 - 156
2 text figs.; 1 table
LoosanorF, Victor Lyon & Frances DoreTTA TOMMERS
1948. Effect of suspended silt and other substances on the rate
of feeding of oysters. Science 107 (2768): 69-70; 2 figs.
NEEDLER, ALFREDA BERKELEY
1934. | Crowding and sex reversal in Ostrea virginica. Ann.
Reprt. Biol. Brd. Canada 1933: 30
THE VELIGER
Vol. 12; No. 1
Orton, JoHn H.
1927. | Observations and experiments on sex-change in the Eu-
ropean oyster (O. edulis). Part I. The change from female to
male. Journ. Mar. Biol. Assoc. 14 (4): 967 - 1045; 4 text
figs.; 12 tables
1933. | Observations and experiments on sex change in the
European oyster (Ostrea edulis). Part III. On the fate of
the unspawned ova. Part IV. On the change from male to
female. Journ. Mar. Biol. Assoc. 19 (1): 1 - 53; 8 text figs.
22 tables
Orton, Joun H. & C. AMIRTHALINGAM
1931. | Observations and experiments on sex-change in the Eu-
ropean oyster (O. edulis). Part II. On the gonad of egg-
spawning individuals. Journ. Mar. Biol. Assoc. 17 (2):
315 - 324; 12 text figs.; 1 table
Rovucu Ley, T. C.
1933. The life history of the Australian oyster Ostrea com-
mercialis, Proc. Linn. Soc. N.S. W. 58 (3/4): 279 - 333;
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ScHaErer, MILNER B.
1938. Preliminary observations on the reproduction of the
Japanese common oyster Ostrea gigas in Quilcene Bay, Wash-
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Sevitta, Maria Luisa H. & EvA Monpracon C.
1965. Desarollo gonadico de Crassostrea virginica GMELIN en
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1: 51-69
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1950. The problem of physiological species with special ref-
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Vol. 12; No. 1
THE VELIGER
Page 37
Levels
of Spontaneous Electrical and Acetylcholinesterase Activities
During Aestivation of the Indian Apple Snail, Pila globosa
BY
P MURALI MOHAN
AND
P MURALI KRISHNA DASS
Department of Zoology, S. V. University College, Tirupati (A.P.), India
(1 Text figure)
INTRODUCTION
Tue INDIAN APPLE SNAIL, Pila globosa (SwaInson, 1822)
aestivates under drought conditions by retiring into the
deeper layers of the mud and sealing itself with the oper-
culum (PrasHap, 1925; Saxena, 1956). The general me-
tabolism of the animal decreases to a low level during
aestivation (MEENAKSHI, 1956; RAaGHUPATIRAMIREDDY,
1965). Under these conditions, it is likely that the nervous
activity could also be low, since the necessity to respond
to external environment does not arise. Moreover, the
altered metabolism in aestivating snails may require a
corresponding altered neuro-regulatory mechanism. Since
the existence of such a mechanism is not known, an at-
tempt was made to study these aspects, taking nervous
activity as well as acetylcholinesterase activity as an index
in aestivated animals.
MATERIALS anp METHODS
Active snails, Pila globosa, were made to aestivate for one
month by embedding them in dry sand in wooden boxes
(Saxena, 1956).
The normal and aestivated Pila were dissected, expo-
sing the different nerve cords under the binocular micro-
scope. The spontaneous electrical activity was recorded by
chlorided silver electrodes. The potentials were fed into
the Tektronix 502A dual beam oscilloscope through Grass
p9 preamplifiers. The photographic recordings were made
using a Grass Kymograph camera.
For the estimation of acetylcholinesterase activity, the
nervous tissue including all ganglia and cords, of normal
and aestivated Pila was isolated and washed repeatedly
with gastropod Ringer solution (HucHES & KERKUT,
1956). The nervous tissue of each animal was used for
a single analysis. The acetylcholinesterase (AChE) activ-
ity was estimated following the modified method of
MetTca tr as suggested by Murat KrisHna Dass (1968).
The incubation mixture contained 0.1 ml of 5% homo-
genate of the nervous tissue in 0.25M sucrose, and 1 ml
of buffer-substrate mixture. After 4 an hour of incubation,
the unreacted acetylcholine was determined by measuring
the optical density at 540 my using Ultraviolet Spectro-
photometer (Hilger and Watts, England), employing
glass cuvettes of 10 mm light path.
RESULTS anp DISCUSSION
The level of spontaneous electrical activity has shown a
definite decrease in all the nerves of the aestivated Pila
from that of the normal as is evident from the recordings
(Figure 1). Both the amplitude and frequency of the
responses were affected on aestivation. A lower level of
spontaneous activity parallels the lowered level of the
active state of the animal (K.P. Rao, 1964; VeNxKaTa-
cHari, 1968). Hence during aestivation, while the animal
remains in a state of torpor, a decreased level of spon-
taneous activity is expected. Similar decrease in electrical
activity was observed in the brain and ganglia of pupae
of diapausing insects (TySHTCHENKO & MANDELSTAM,
1965; Kutyna & TomBes, 1966).
40onV
|
0.4. sec
Figure 1
Spontaneous activity recorded from cerebral commissure (a1, a2),
supra-intestinal nerve (b 1, b2) and left pleuro-visceral connective
(c 1, c2) in normal (a1, b1, c1) and aestivated (a2, b2, c 2)
Pila globosa
Note the decrease in the spike amplitude and frequency of pulses
on aestivation
A direct relationship between electrical activity and
AChE activity was demonstrated in relation to the behav-
ior of scorpion (VENKATACHARI & Murat KrisHNA
Dass, 1968). Since in the present case a considerable
decrease in the spontaneous activity was observed, it is
possible that the associated AChE activity might also
have a corresponding decrease in activity. As is evident
from ‘Table 1, there is a decrease in the AChE activity level
Table 1
Levels of acetylcholinesterase enzyme activity in normal
and aestivating Pila globosa. (Activity is expressed in mg
of acetylcholine metabolised per gm weight of tissue
per hour).
Normal snails Aestivating snails % decrease
during
aestivation
338 + 20.09 281.33 + 17.78 16.76
(P > 0.001)
Each value in this table is an average of six separate
analyses (six animals).-
THE VELIGER
Vol. 12; No. 1
in nervous tissue of aestivated snails. This observation may
possibly reiterate the direct relationship between elect-
rical activity and AChE activity.
Since the nervous system in general is known to regulate
the activities of the animal, the lowered nervous activity
of aestivating snails in terms of decreased spontaneous
and acetylcholinesterase activities could be responsible for
their observed lowered metabolism. Further work along
these lines is in progress.
SUMMARY
1. The spontaneous electrical and acetylcholinesterase ac-
tivities were studied in the nervous tissue of normal
and aestivated Pila globosa.
2. The spontaneous activity decreased in aestivated snails.
Correspondingly, there was a decrease in the AChE
activity.
ACKNOWLEDGMENTS
The authors wish to express their thanks to Dr. K.S.
Babu and Dr. K.S. Swami of this Department for their
kind help and valuable suggestions.
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THE VELIGER
Pampapati, Rao K.
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Page 39
Page 40
THE VELIGER
Vol. 12; No. 1
Macoma (Psammacoma) pulleyi, a New Clam from Louisiana
BY
PAUL S. BOYER
Department of Geology, Rice University, Houston, Texas ‘77001
(2 Text figures)
THE FOLLOWING PELECYPOD was dredged by the author
in the course of investigations of invertebrate ecology off
the coast of Louisiana. Reference to published descrip-
tions and comparison with the collections in the U.S.
National Museum indicate that this clam is a new species.
PELECYPODA
TELLINIDAE
Macoma Leacu, 1819
(Psammacoma) Dati 1900
Macoma (Psammacoma) pulleyi Boyer, spec. nov.
(Figure 1)
Description: Shell thin, white, moderately inflated, elon-
gate, inequilateral, equivalve, and covered by a very thin,
brown periostracum. Anterior end longer, the anterior
dorsal margin almost rectilinear, the anterior margin
rounded above and more gently and evenly curved below.
Ventral margin nearly straight, subparallel to the anterior
dorsal margin, and intersecting the posterior margin ab-
ruptly at an angle of about 120°. Posterior dorsal margin
sloping steeply, and meeting the posterior margin in a
curve. Surface of shell beneath periostracum lusterless
white, and smooth except for very fine incremental growth
lines; an obtuse, rounded ridge running from the umbo
to the posterior ventral angle marks the intersection of
the posterior slope with the surface of the disk: this ridge
is better defined in the left valve, where the posterior
slope is occupied by a broad, very shallow sulcus. Hinge
plate very narrow, bearing in the left valve a bifid ante-
rior cardinal tooth and a lamellar posterior cardinal tooth;
right valve with a bifid anterior cardinal tooth and a
smaller, grooved posterior cardinal tooth. Ligament area
shallow; posterior area not sharply defined. Pallial sinus
mildly discrepant between the valves: in the left valve,
obliquely rising from the posterior adductor muscle scar,
sinuous above, narrowing and extending forward about
= of the distance between the adductor muscle scars,
bluntly rounded anteriorly, about half confluent with the
pallial line below; in the right valve obliquely rising from
the posterior adductor muscle scar, smoothly rounded
above, sloping in a straight line anteriorly, somewhat less
bluntly rounded anteriorly, extending forward about # of
the distance between the adductor muscle scars, about half
confluent with the pallial line below. There is a slight
posterior gape (about 1.5 mm in the holotype), and the
posterior end is angled to the right almost imperceptibly.
The anterior dorsal margin of the right valve overlaps
the left valve slightly; in front of the umbo of the left
valve there is a small incised area for the partial recep-
tion of the overlapping portion of the right valve.
Measurements (in millimeters)
Length Height Diameter
Holotype 42.0 Done 12.4
1* Paratype 40.8 opacd) 12.0
2¢ Paratype 44.5 23.8 13.9
3™ Paratype _ 28.6 15.1 8.3
4" Paratype 55.3 29.8 16.7
Type Locality: The holotype was collected live offshore
from Plaquemines Parish, Louisiana, 28°59’ N Latitude,
89°30’ W Longitude, 18 m water depth, in silt substrate,
salinity (interpolated from adjacent stations) 15%, at
surface, 31%, at bottom, 9 June 1965.
Known Range and Habitat: Macoma pulley: was collec-
ted west of the Mississippi Delta, offshore from Terre-
bonne, Lafourche, Jefferson, and Plaquemines parishes,
Louisiana, in mud substrate, in water depths of from
Vol. 12; No. 1
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Page 41
13 to 45 m, in bottom water having nearly normal salin-
ity (30 to 33%). The unusually large 4" paratype was
dredged dead in 19m water depth farther west, from a
patch of sandy mud substrate 52 km south of the mouth
of the Mermentau River, Cameron Parish, Louisiana.
Parker illustrates this species (PARKER, 1956; plt. 1,
figs. 8A, 8B; the same pictures appear in Parxer, 1960,
pit. 4, figs. 14A, 14B) as Macoma tageliformis. He lists
several localities (PARKER, 1956, p. 315, table II) for M.
tageliformis which probably refer to M. pulleyi. These
localities are east of the Mississippi Delta: live specimens
were common at the delta front, and dead valves were
found in the shallow shelf area.
United States National Museum no. 189186, collected
from Louisiana by L. R. Cary, is this species.
Disposition of Specimens: Holotype deposited in the Mu
seum of Comparative Zoology at Harvard University, no.
271577; 1* paratype (Figure 1) in the author’s collection;
2™ paratype in the U. S. National Museum; 3°: paratype
Figure 1
Macoma pulley Boyer, spec. nov.
First Paratype
A. Interior, right valve
C. Exterior, right valve
— matching valves
B. Interior, left valve
D. Exterior, left valve
in the Academy of Natural Sciences of Philadelphia;
4" paratype in the Bureau of Economic Geology, The
University of Texas, Austin.
Remarks: Macoma (Psammacoma) pulleyi has been con-
fused with the sympatric M. (P) tageliformis Dat, 1900.
Macoma tageliformis was originally described from Cor-
pus Christi Bay, Texas, but was not figured (Dau, 1900a,
pp. 300, 315). Another description, also without figures,
appeared about a month later (Dat, 1900b, p. 1055).
Subsequently, illustrations were published of a specimen
from Puerto Rico (Dati « Simpson, 1901, plt. 55, figs.
10, 11, 15) which was identified as M. tageliformis. Dati
« Smmpson’s figured specimen (U.S. National Museum
no. 160497) is relatively much shorter than the holotype
of M. tageliformis (U.S. National Museum no. 6086) ;
the illustrated specimen also differs from the holotype in
having a steeper posterior dorsal margin, which makes it
look superficially like M. pulley:. Thus Datw’s choice of
specimen to illustrate as Macoma tageliformis may have
caused some of the confusion surrounding that species.
Collections of Macoma tageliformis from Louisiana
agree perfectly with Datt’s holotype. Figure 2 is a speci-
men of M. tageliformis from sandy silt substrate, 19 m
water depth, 39km south of Rollover Bayou, offshore
from Vermilion Parish, Louisiana.
Macoma tageliformis and M. pulleyi may easily be
distinguished by the following differences:
a) Macoma tageliformis is more noticeably inequivalve,
its left valve being more convex than its right.
b) In Macoma tageliform: ‘he ligament area is depressed,
so that the surface of .he posterior area on either
side is almost vertical and is set off from the rest of
the shell by a rather sharp ridge; the posterior area
is not so defined in Macoma pulleyi.
M oma pulley: has a rather abrupt posterior ventral
an ,ie; also
Macoma pulley: bears a ridge delineating the poste-
rior slope from the surface of the disk. These last two
features are lacking in M. tageliformis.
Figure 2
Macoma tageliformis Dat, 1900
Matching valves. Offshore from Vermilion Parish, Louisiana
A. Interior, right valve B. Interior, left valve
C. Exterior, right valve D. Exterior, left valve
Page 42
e) The pallial sinus of Macoma pulleyi extends farther
forward than that of MM. tageliformis.
The closest relative of Mfacoma pulleyi would seem to
be the Miocene species M. holmesii Dati, 1900, from
Duplin County, North Carolina.
Macoma pulleyi is dedicated to my friend Dr. T. E.
Pulley, of the Houston Museum of Natural Science. Dr.
Pulley is an authority on the mollusks of the Gulf Coast.
ACKNOWLEDGMENTS
The author wishes to acknowledge the support of a
National Aeronautics and Space Administration fellow-
ship and of National Science Foundation Grant GP-3600,
and the use of the facilities of the Geology Department
of Rice University. Dr. Robert R. Lankford supervised
the project of which this study is a part; he also critically
read the manuscript and contributed many helpful sug-
gestions.
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Vol. 12; No. 1
LITERATURE CITED
Dati, WiLL1iAM HEALEY
1900a. Synopsis of the family Tellinidae and of the North
American species. Proc. U.S. Nat. Mus. 23 (1210) : 285 to
326; plts. 2-4 (14 November 1900)
1900b. Contributions to the Tertiary fauna of Florida, with
especial reference to the silex-beds of Tampa and the Pliocene
beds of the Caloosahatchie River, including in many cases a
complete revision of the generic groups treated of and their
American species. Part V. Teleodesmacea: Solen to Diplodon-
ta. Trans. Wagner Free Inst. Sci. Philadelphia 3 (5) : 949 to
1218; plts. 36 - 47 (December 1900)
Dai, WiLtiaM HEALEY & CHARLES TorrEY SIMPSON
1901. The Mollusca of Porto Rico. U.S. Fish Comm.
Bull. 20 (1): 351-524; plts. 53 - 58 (November 1901)
Parker, Rospert H. ;
1956. Macro-invertebrate assemblages as indicators of sedi-
mentary environments in east Mississippi Delta region.
Bull. Amer. Assoc. Petrol. Geol. 40 (2): 295-376; 32 figs.;
8 plts.
1960. Ecology and distributional patterns of marine macro-
invertebrates, northern Gulf of Mexico. In: Recent sediments,
northwestern Gulf of Mexico, ed. F R. SHeparp, EB. PHLEGER
« T.H. Van ANDEL, pp. 302 - 337; 6 plts.; 17 text figs.
Amer. Assoc. Petrol. Geol., Tulsa, Oklahoma
Vol. 12; No. 1
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Page 43
On Pseudopythina rugifera (CARPENTER, 1864) (Bivalvia)
WALTER NARCHI
Faculdade de Filosofia, Ciéncias e Letras da Universidade de Sao Paulo, Sao Paulo, Brasil
(9 Text figures)
INTRODUCTION
LitTLE Is KNOWN of the structure, functions of the organs
in the mantle cavity, and the adaptations found in the
Leptonacea, composed largely of commensal species, in
correlation with their specialized mode of life.
PopHaM (1940, p. 549) refers to the biology of some
species belonging to the genera Kellia, Lasaea, Montacuta,
Mysella, Devonia and Galeomma. Ceratobornia cema
(= Lepton cema Narcut, 1966) was intensively studied.
Other genera were studied but no references were found
for Pseudopythina.
Pseudopythina rugifera (CARPENTER, 1864) occurs on
the Pacific Coast of North America from Alaska to Lower
California (ApBoTT, 1954, p. 395; PALMER, 1958, p. 90).
According to Kren (1937, p. 25) it ranges from 37° to
48° North Latitude. The species is well known in the
eastern Pacific as an “ectoparasite” of the crustacean
Upogebia pugettensis (Dana, 1852) and could be found
attached by its byssus to the ventral surface of the poly-
chaete worm, Aphrodita (OLpRoyp, 1924, p. 136; Mac-
Ginitre & MacGrinrrig, 1949, p. 348; Parmer, loc. cit.;
Boss, 1965, p. 186). Otproyp (Joc. cit.) states that all
species of the genus Pseudopythina FiscHER, 1884, are
commensals. In this paper observations on the structure,
ciliary currents of feeding and digestion, and some other
functional adaptations of P rugifera are studied.
The animals were found attached to the broad annu-
lated foot of Aphrodita refulgida Moore, collected at
Tomales and at Bodega Bay, Marin County, California
(Figure 1). Several specimens were examined alive and
the ciliary currents in the mantle cavity were observed
with use of carmine, aquadag, and powdered carborun-
dum. Others were fixed in Bouin’s fluid. Serial sections
of 6m were made and the sections stained in Delafield’s
hematoxylin, eosin and Alcian blue, to examine the gen-
eral anatomy. The observations on living specimens were
made at the Pacific Marine Station, Dillon Beach, Cali-
fornia.
Figure 1
General view of several specimens of Pseudopythina rugifera
attached to Aphrodita
SHELL anp MANTLE
The shell is small, oval-oblong, moderately obese and frag-
ile, reflecting the protected habitat of this species. The
umbones are close together, and located near the middle
line of the shell. Lateral teeth are absent and one large
central tooth (Figure 2) exists on each valve (CarPEN-
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Vol. 12; No. 1
Page4$ 0 Oe
Figure 2
View of the hinge of Pseudopythina rugifera (X 40)
TER, cited by PaLmeEr, 1958, p. 90). The shell is covered
with a thin, light brown periostracum which is concent-
rically wrinkled. The embryonic shell is still visible in the
adult near the umbo. Foraminifera were found frequent-
ly attached to the shell. The shell of the largest animal
encountered in this study measured 6mm in length,
2.5 mm in height, and 1.5mm in width.
The species possesses both an external and an internal
ligament, the latter being much more developed. As in
Kellia suborbicularis (Montacu, 1803), the shell and
the fleshy parts of Pseudopythina rugifera are white in
color (OtprreLp, 1961, p. 257). In living animals the
foot protrudes from the shell to a distance equal to half
the length of the shell. The mantle edges, the inhalant
and exhalant siphons do not extend beyond the shell valves
as in Montacuta substriata (PopHam, 1964, p. 564).
As in Montacuta ferruginosa and M. substriata (OLp-
FIELD, 1961, p. 260), there are only two pallial openings:
a large, inhalant and pedal aperture in the anterior and
ventral part of the body, and a small exhalant aperture
posteriorly located. The inner mantle folds are fused be-
tween these two apertures in a very short extension.
The exhalant siphon is shorter and formed by the
inner mantle folds. The edges of this siphon bear small
papillae of which one dorsal and one ventral papilla are
slightly larger and can be seen protruding out of the shell.
The mantle edge has three folds. The outer fold is
enclosed by the periostracum and has a flattened epi-
thelium. The middle fold bears few papillae and secretes
the periostracum. The inner fold contains the circumpal-
lial nerve. The folds are very similar to those of Kellia
suborbicularis (OLDFIELD, 1961, p. 260). In the dorsal
part of the inner mantle fold, from the region of the
stomach up to the pallial fusion, a tract of strongly cili-
ated cells occurs. This tract is concerned with the rejection
of particles from the mantle. The free edge of the mantle
surrounding the inhalant siphon and pedal opening is
also ciliated. Particles falling on this edge are passed into
the mantle cavity. ,
Tue CTENIDIA
Both demibranchs are present in Pseudopythina rugifera
(Figure 3), but the outer demibranch (od) is less than
half the depth of the inner (id). Only the inner demi-
branch has a food groove along its ventral edge. Lam-
mellae are flat and homorhabdic, as found in the Leptoni-
dae (Atkrns, 1937, p. 391). Behind the foot (f), the
left and right inner demibranchs are joined together in
the median plane. As in Kellia suborbicularis (OLDFIELD,
1961, p. 263), both outer and inner demibranchs on each
side are fused ventrally to the mantle, in the region where
the fused inner folds of the mantle separate the common
inhalant and pedal aperture from the exhalant aperture.
Hin
Figure 3
Pseudopythina rugifera (CARPENTER, 1864)
Mantle cavity viewed from the left side after removal of the left
shell valve and mantle lobe, showing ciliary currents.
a-— anus am — anterior adductor muscle _ b — byssus
arp — anterior retractor pedis muscle dd — digestive diverticula
exs — exhalant siphon f — foot g — foot groove
id — descending lamella of inner demibranch __ li—inner labial palp
lo—outer labial palp | oa—ascending lamella of outer demibranch
prp — posterior retractor pedis muscle | w — waste material
pm — posterior adductor muscle te — tentacle
The gill filaments are numerous. Slender interfilamentar
junctions are numerous but interlamellar junctions are
few. This condition is found in other species where the
incubatory habit occurs (OLDFIELD, 1961, p. 263).
The ciliary mechanism in the ctenidia of Pseudopythina
rugifera is similar to that of Kellia suborbicularis (At-
Vol. 12; No. 1
KINS, 1937, p. 389). The ctenidia and their ciliation are
as in Type C(1) of Atkins (loc. cit.).
There is no interruption of the latero-frontal cilia at
the ventral tip of the filaments of the ctenidia, but rather
a bending of the filaments. The frontal currents on the
outer demibranch are directed ventrally on the ascending
lamella, bend at the free edge and flow dorsally on the
descending lamella. The frontal cilia are not modified. In
the free edge of the outer demibranch no longitudinal
currents occur.
The frontal currents on the inner demibranch are direc-
ted ventrally on the ascending and descending lamellae.
In the free edge of the inner demibranch an oral longi-
tudinal current occurs. Other oral currents occur between
the bases of the two demibranchs of each side of the
body.
Along the marginal grooves of the inner demibranch
shorter guarding cilia exist. On each ventral tip of the
filaments these cilia beat as a group, moving particles to
the anterior part of the body.
The fine frontal cilia are continued on each side and
beat directly to the ventral part.
eT
EAI WEN
oie tide
OH %
a BN
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Page 45
The long lateral cilia form an inhalant current and the
frontal cilia convey food particles along the filament to
the food groove.
There is one type of latero-frontal cilia: the eu-latero-
frontal cilia which are fused, large, and prevent the loss
of food particles into the supra branchial chamber.
As in Kellia suborbicularis (OLDFIELD, 1961, p. 264),
guarding cilia exist on both sides of the food groove and
these cilia prevent the entrance of coarse particles into
the groove. ATKINS (1937, p. 360) suggests that guarding
cilia occur when the animals live in a silt or muddy sub-
strate.
MUSCULATURE
The anterior adductor muscle (Figure 4, am) is slightly
larger than the posterior adductor muscle (pm). The re-
jected material is violently shot out of the pallial cavity
by a sudden contraction of the adductors.
PopHam (1940, p. 564) describes the same phenomenon
for Kellia suborbicularis. OLDFIELD (1961, p. 264) believes
Figure 4
Pseudopythina rugifera (CARPENTER, 1864)
Diagrammatic representation of the organs in the mantle cavity
am — anterior adductor muscle
arp — anterior retractor pedis muscle
bl — byssogenous lamella
dd — digestive diverticula
i — intestine
au — auricle
bgi- main byssus gland
cg — cerebral ganglion
f — foot g — foot groove
ia — ascending lamella of inner demibranch
ss — style sac
oa — ascending lamella of outer demibranch
prp — posterior retractor pedis muscle
ve — ventricle
id — descending lamella of inner demibranch
li — inner labial palp _lo — outer labial palp
k — kidney
bv — ovary
oe — oesophagus
pm — posterior adductor muscle
r—rectum s—stomach
pg — pedal ganglion
vg — ventral ganglion
Page 46
that this is probably essential in species which live in a
sandy or muddy habitat and prevents the danger of
their silting up. Each adductor muscle is subdivided equal-
ly into two parts, as in Kellia and Montacuta.
There are two pairs of muscles of the foot: a pair of
anterior retractor pedis muscles (arp) and a pair of pos-
terior retractor pedis muscles (prp).
As in Kellia and Montacuta, there is no elevator pedis
muscle. The anterior retractor pedis muscle is completely
subdivided into a smaller retractor pedis muscle and a
larger byssus retractor muscle. As in Montacuta substriata
(OLDFIELD, 1961, p. 267) the well developed byssus mus-
culature is related to the high development of the byssus
apparatus in this species. In Pseudopythina rugifera the
protractor pedis muscle runs through the lower part of
the anterior retractor pedis muscle, but is inserted to-
gether with the byssus retractor muscle (br), while in
M. substriata they have a separate origin from the shell
(OLpFIELD, 1961, p. 267).
THe FOOT
The foot is large and though laterally compressed, has a
flat creeping sole by means of which it is capable of
active locomotion. It is slender and, when fully distended,
is half the length of the shell, but it can be completely
withdrawn when the shell valves are closed. The large
pedal aperture combined with the inhalant aperture per-
mits the foot to move in a wide angle. As in Montacuta
substriata (PopHam, 1940, p. 565), rocking movements
are possible because of the gape and the large size of the
pedal opening. The byssus cavity (bg) lies posteriorly in
the foot and opens by way of a byssus canal into a groove
(g) which extends on the ventral side almost to the tip
of the foot. In M. substriata (PopHaM, loc. cit.) two or
three threads are produced but in Pythina rugifera the
number is greater.
The ventral and ventro-lateral surfaces of the foot
are covered by a ciliated columnar epithelium, as are the
surface of the byssogenous lamellae (bl), the byssus
canal and groove.
The viscera do not extend into the foot. In the foot there
are many muscle fibers and connective tissue with large
blood spaces.
Pseudopythina rugifera lives attached to the foot of
the sea mouse by a large byssus (b) composed of a large
number of threads. The byssus cavity is embedded in the
main byssus gland and is divided by a fold into two large
longitudinal parts (Figure 5). Each part is divided into
10 to 12 slit-like compartments by the byssogenous lamel-
lae. Each lamella is composed of connective tissue and
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Vol. 12; No. 1
Figure 5
Pseudopythina rugifera (CARPENTER, 1864)
Byssus gland and byssus thread
fine muscle fibers from the byssus retractor muscle. A
ciliated columnar epithelium covers the lamellae and is
composed of low columnar cells. The byssus apparatus of
P. rugifera is very similar to that found in Potidoma sub-
trigonum (Derrovux, 1961, p. 119).
The glandular cells of the main byssus gland (bgi)
surround the byssus cavity. From these cells, in which the
cytoplasm laden with fine granules, stains in aniline blue,
long slender ducts lead to openings between the ciliated
epithelial cells covering the surfaces of the byssogenous
lamellae, in the byssus cavity. Thus the secretion of the
main byssus gland forms a coating to the lamellae, and
these sheets of secretion coalesce to form the stalk of the
byssus.
The byssus canal is ciliated on its anterior and lateral
walls and the byssus groove is ciliated throughout its
entire length.
Vol. 12; No. 1
THE VELIGER
Page 47
eee
The main mucus gland occupies the anterior part of
the foot and extends along both sides to the posterior
end of the foot. From these mucus cells slender ducts open
on the byssus groove on the ventral and ventro-lateral
surfaces of the foot.
The construction of the byssus threads by adult animals
was observed in the laboratory. A new byssus thread is
constructed in 30 seconds and this velocity is probably
due to the habits of the animal. To build a new byssus
thread the foot is placed on the sole of the sea mouse. It
becomes temporarily attached, apparently by a sucker-
like action, and does not move. Only the tip of the foot
moves laterally back and forth, until the byssus thread is
completed.
Pseudopythina rugifera, detached from the sole of Aph-
rodita, with very active locomotion attached itself to the
ventral part of the animal in a few seconds. As in Mont-
acuta substriata (PopHaM, 1940, p. 566), the surface of
the foot is strongly ciliated. Particles move rapidly across
the sides of the foot from the tip to the heel and are
passed off into the main rejection current concerned with
the rejection of waste material, which passes out between
the shell valves in a posterior and ventral direction im-
mediately behind the foot.
LABIAL PALPS
The labial! palps are relatively small; their opposed sur-
faces are covered by cilia and they have 9 transverse
ridges. The ridged and grooved surfaces function as a
sorting region. Particles of food and other material are
swept onto the palps from the food groove of the inner
demibranch. On the crests ot the folds, particles are
carried forward from fold to fold towards the mouth. On
the floor of the grooves, between adjacent folds, particles
are driven to the ventral border of the palp and from
there moved to the tip of the palp where they are rejected.
The ventral tips of the anterior filaments of the inner
demibranch are not inserted into a distal oral groove,
although the antero-ventral margin of the inner margin
of the inner demibranch is fused to the inner palp lamella
and belong to the Category III of the association of
ctenidia and labial palps (Stasex, 1963, p. 91).
THe ALIMENTARY CANAL
GENERAL STRUCTURE
The small size of the specimens made it difficult to
study and observe the internal structure of the stomach
in great detail. The alimentary canal was dissected in
specimens that had been preserved in alcohol. The in-
ternal structure of the stomach was studied in living
specimens. The stomach was opened from the surface by
a mid-dorsal incision through the roof, and the right side
of the stomach was drawn downwards. The ciliary cur-
rents were determined with the aid of carmine, aquadag
and fine carborundum particles. The nomenclature used
by GraHam (1949), Owen (1953), PurcHoN (1955),
and Rem (1965) has been followed.
The mouth opens into the oesophagus ( oe) which joins
the stomach (s) at the anterior and ventral part, which
is thin-walled and globular. The stomach is approximately
cylindrical in shape and is surrounded by the digestive
diverticula (dd).
The combined style-sac (ss) and intestine open into the
posterior and ventral region of the stomach; the style-sac
and intestine intercommunicate by an opening. This open-
ing was observed in Montacuta by PELSENEER (cited by
OpriELp, 1961, p. 273) but it was not seen by her in
the same genus. The style-sac and intestine are joined in
a short extension; after this, the intestine leaves the style-
sac on the right side. The intestine (i), after a small loop,
ascends dorsally and then continues as a very well de-
veloped rectum (r).'The rectum passes over the posterior
adductor muscle and finally opens into the anus (a) close
to the exhalant aperture.
STRUCTURE or tut STOMACH
Pseudopythina rugifera has a stomach of type 4 (Figure
6), as defined by Purcuon (1958, p. 488). The minor
typhlosole is absent in this species. PuRCHON (op. cit., p.
489) states that the minor typhlosole appears to be ab-
sent in the Anomiidae, Erycinidae and Montacutidae.
The major typhlosole (ty) projects into the stomach, pas-
ses forwards from the aperture of the mid-gut and
curves gradually to the left over the floor of the stomach.
The major typhlosole is accompanied on its right side by
the intestinal groove (ig).
A sorting area consisting of a small number of folds
and grooves is present on the anterior face of the stomach
between the orifice of the oesophagus (oe) and the intes-
tinal groove. This sorting area extends on to the left
anterior wall of the stomach, invades the left pouch (Ip)
and ends close to the terminal region of the major typhlo-
sole.
The intestinal groove is enveloped by the sorting area.
There is a relatively small dorsal hood (dh), the aper-
ture of which is penetrated by a lobe of the gastric shield
(Figure 7, gs). The dorsal hood lies on the left anterior
Page 48
Figure 6
Pseudopythina rugifera (CARPENTER, 1864)
Interior of the stomach, seen from the right side, after an incision
through its right side extending into the style sac
dh - dorsal hood gs — gastric shield
Ip — left pouch mg — midgut
oe — oesophagus ss — style sac
ig — intestinal groove
r, — ciliated ridge
ty — major typhlosole
Figure 7
Pseudopythina rugifera (CARPENTER, 1864)
Transverse section of the stomach
ep — epithelium gs — gastric shield _st — style
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Vol. 12; No. 1
stomach wall, as in Scintilla hanleyi (PuRcHOoN, 1958,
p. 498).
The epithelium of the stomach underlying the gastric
shield consists of tall narrow columnar cells each with an
oval nucleus near the base of the cell (Figure 8). The
distal region is packed with almost spherical granules up
to 4u in diameter. Similar spherical granules were found
in cells of the stomach of Nuculidae (Owen, 1953, p.
545). Surrounding the cells of the epithelium there is a
layer of collagen and a system of smooth circular muscle
fibers.
The crystalline style projects obliquely upwards and its
tip is placed in the aperture of the dorsal hood.
The left pouch is on the left wall of the stomach. The
upper border of the left pouch is penetrated by a lobe
Aen antes wa ih Wy
Lali tit
40h
Figure 8
Epithelium of the stomach showing spherical granules
of the gastric shield. The left pouch receives one duct
from the digestive diverticula on the left side of the body.
In the grooves of the sorting area the cilia beat to-
wards the intestinal groove, while the cilia on the tops
of the ridges beat towards the dorsal hood as in Gale-
omma glycymeris (Reip, 1965, p. 169).
A circulating current is set up by the cilia on the top
of the sorting area. It is this general circulation which
Vol. 12; No. 1
carries large particles and masses of particles towards the
dorsal hood and gastric shield. Light particles are carried
in the current flowing across the ridges of the sorting
area into the dorsal hood. Heavy particles are driven
against the surface of the sorting area and fall into the
transverse rejection grooves. Small and heavy particles
are carried away to the intestinal groove and then into
the mid-gut.
The sorting area in Pseudopythina rugifera is composed
of large ridges and wide grooves. The cilia on the crests
of the folds beat, carrying particles to the left region of
the stomach. As in Scintilla hanley: (PuRcHoN, 1958, p.
499) there is no sorting area within the dorsal hood, nor
is there one on the right wall of the stomach.
From the oesophagus a narrow ciliated ridge (r:) passes
backwards from the left corner of the oesophageal orifice
to the dorsal hood. The dorsal hood receives particles from
the anterior part of the stomach by means of that ridge.
A similar ridge was described by PurcHon (1958, p.
499) in Scintilla hanleyi.
The sorting area is small; large particles are not rejec-
ted and excessive quantities of small particles are accepted
(PurcHon 1958, p. 487).
EXCRETORY SYSTEM
The kidney (Figures 9 C, 9 K) is composed of a pair of
elongated sacs with glandular walls which intercommuni-
cate in the median part, ventral to the pericardial cavity.
Each kidney sac is lobed ventrally and dorsally and there
are no internal folds of the wall to produce spongy kid-
ney. It opens directly at its anterior end to the supra-
branchial chamber by the external renal aperture, which
has ciliated cells. These ducts (rpd) are long and they run
posteriorly from the pericardial cavity and open into the
posterior part of the kidney. Histologically this system is
very similar to that of Kellia and Montacuta (OLDFIELD,
1961, p. 276).
NERVOUS SYSTEM
There are three pairs of ganglia: the cerebral, pedal,
and visceral. The cerebral ganglia lie behind the anterior
adductor muscle, one on each side of the oesophagus,
and joined together by a supra-oesophageal commissure.
The buccal nerve, the anterior adductor nerve, and the
anterior retractor pedis muscle nerve arise from these
ganglia.
The pedal ganglia are so close together that they ap-
pear as one, lying in the mid-line ventrally to the digestive
THE VELIGER
Page 49
diverticula. Each pedal ganglion gives rise to the anterior
pedis nerve, the lateral pedis nerve, and the byssus nerve.
The visceral ganglia lie on the anterior surface of the
posterior adductor muscle, and have the branchial nerve,
the posterior adductor nerve, and a posterior pallial nerve.
The cerebro-visceral connective and the cerebro-pedal
connective are similar to those described by OLpFIELD
(1961, p. 280) for the genera Kellia and Montacuta.
REPRODUCTIVE SYSTEM
The specimens examined were females. The gonad is a
large hollow organ with two anterior lobes, each subdi-
vided into many lobules and communicating with a single,
median posterior chamber. As in Kellia suborbicularis
(OLpFIELD, 1961, p. 281), the posterior chamber opens
on each side of the body by a short ciliated duct into the
supra-branchial chamber of the inner demibranch. The
external genital opening is immediately anterior to the
exterior renal opening.
As in Kellia suborbicularis (LEsour, 1938, p. 447) and
other specialized eulamellibranchs (OxpFIELD, 1964, p.
79) fertilization occurs probably within the gills, the sper-
matozoa of another individual being drawn in with the
inhalant current.
The specimens collected in April contained developing
embryos (e) within the gills, and the gonad appeared
empty with a few small atretic ova. Pseudopythina rugi-
fera is not a common animal in Tomales Bay and further
investigations may reveal male or hermaphrodite forms.
More extensive collecting of animals would reveal if the
species is a protogynous consecutive hermaphrodite in
which there is no overlapping of male and female phases,
as in Kellia suborbicularis and Montacuta ferruginosa
(OLpFIELD, 1961, p. 289).
DISCUSSION
Pseudopythina rugifera is adapted to live attached and is
found fixed by its byssus to the ventral surface of Aphro-
dita or Upogebia. The species attaches and detaches itself
at will, as observed by MacGrniTiE and cited by QUAYLE
(1960, p. 74). The animal has a well developed foot and
a wide pedal gape to allow the foot to be extruded. It
has two pallial apertures, a large antero-ventral and a
small posterior one.
Concerning the ciliary mechanism of the ctenidia the
species is similar to Kellia suborbicularis (OLDFIELD, 1961,
p. 264) as both have guarding cilia in both sides of the
food groove on the inner ctenidia. This suggests that the
Page 50
(adjacent column —>)
animal lives in a silt or muddy substrate. The manner in
which the rejected material is violently shot out of the
pallial cavity by a sudden contraction of the adductors
proves that the animal lives in a sandy or muddy environ-
ment (OLDFIELD, loc. cit.).
The byssus apparatus is well developed with a large
byssus retractor muscle as in Montacuta substriata which
lives attached to the spines of the sea urchins. The byssus
apparatus is divided in the middle by a fold into two longi-
tudinal parts. Each part is divided into 10 to 12 slitlike
compartments and is similar to Potidoma subtrigonum
(JEFFREYS) described by Deroux (1961, p. 119). It is
different from the other species belonging to the genera
Kellia, Montacuta, and Lasaea.
The absence of a well developed sorting area in the
stomach suggests that excessive quantities of small par-
ticles or large particles are not rejected.
The rectum is very wide. The contents of the intestine
THE VELIGER
Vol. 12; No. 1
SS
OY
oe S|
> tom, 8
40 eS . Y
* ‘ —s Q
ri
aS e
a \* \
Al ©
=. oe ss
ov # fs ANS Sass) 8
| z b A s g.Os
il. ° 6 Py SS
Spe i GS
fo fg
aaa
Wa snmandes Y
bgi Ley
WN ff 4
i VV 4
ie: =f
mil
mml
mol
Figure 9
Pseudopythina rugifera (CARPENTER, 1864)
Diagrammatic transverse sections through the animal (X 72)
A — at level of the stomach B — at level of the style sac
C — at level of the kidneys
bg — byssus cavity bgi — main byssus gland
br — byssus retractor muscle dd — digestive diverticula
e — embryos ex! — external ligament fg — food groove
g — foot groove gs — gastric shield _i — intestine
id — descending lamella of inner demibranch __ il—internal ligament
k — kidney _mil — inner fold of mantle edge
mml — middle fold of mantle edge od — outer demibranch
mol — outer fold of mantle edge _— ov — ovary
prp — posterior retractor pedis muscle r — rectum
rpd — reno-pericardial duct _ ss — style sac _st — style
when compared with those of the stomach show a larger
number of more or less complete but empty diatom tests
and protozoan skeletons. This suggests that while some
Vol. 12; No. 1
lo)
OQ
|
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00
for
sf :
Ss a 5 %
ss yo
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Oo 8 8 C os Sg -
See gWVZ ghee: iss”
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EPISODIO A ds
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(<— on facing page)
mechanical trituration occurs in the stomach, enzymes
are also present (OWEN, 1956, p. 559).
The stomach of Pseudopythina rugifera is in some
aspects similar to that of Scintilla hanleyi, as both have
two ciliated ridges from the anterior left face of the
stomach to the dorsal hood. The oesophagus, the left
pouch and the dorsal hood are similar in both species.
While in P rugifera the style sac and the intestine inter-
communicate by an opening, this is not found in S. hanley:.
The style sac and intestine join in a short extension and
the intestine leaves the style sac laterally on the right
side.
The sorting area in Pseudopythina rugifera starts in the
right wall of the stomach, passes upwards from the region
of the intestinal groove towards the oesophageal aperture
to the left wall of the stomach and then surrounds the
left pouch, ending close to the major typhlosole. Scintilla
THE VELIGER
Page 51
hanleyi has two sorting areas (PuRCHON, 1958, p. 498) :
S; on the anterior face of the stomach which passes from
the intestinal groove to the oesophageal aperture, and Su,
the second sorting area, on the floor of the left pouch.
It seems that P rugifera has a stomach more primitive
than S. hanleyi, as the two sorting areas are joined into
only one in the present species.
The stomach of Pseudopythina rugifera is an example
of simplification of structure: the major typhlosole is
reduced. The sorting area directs particles toward the
intestinal groove.
The stomach of Pseudopythina rugifera probably tri-
turates ingested particles by muscular action and the rela-
tively large size of the stomach and style sac is an adapt-
ation for the ingestion of large quantities of bottom ma-
terial (OwEN, 1956, p. 562).
SUMMARY
Pseudopythina rugifera (CARPENTER, 1864) lives attached
to the ventral surface of the polychaete worm Aphrodita
and to the anomuran crustacean Upogebia. It occurs on
the Pacific Coast of North America, from Alaska to
Lower California. The shell is small, oval-oblong, and frag-
ile, reflecting the protected habitat of the species. There
are only two pallial apertures, a large anterior and vent-
rally located one which is a common inhalant and pedal
opening, and a small exhalant aperture. The siphons do
not protrude beyond the shell.
The ctenidia are similar to those of Kellia suborbicularis
and both outer and inner demibranch on each side are
fused ventrally.
The byssus apparatus is well developed and is very sim-
ilar to that of Potidoma subtrigona (DERoux, 1961, p.
1).
The anatomy of the stomach is described in detail, and
the stomach belongs to Type 4 (PurcHon, 1958, p. 488).
Specimens collected in April contained developing em-
bryos within the gills. Future research will prove if Pseu-
dopythina rugifera is a protogynous consecutive herm-
aphrodite in which there is no overlapping of male and fe-
male phases as in Kellia suborbicularis and Montacuta
ferruginosa.
ACKNOWLEDGMENTS
I wish to express my thanks to Dr. Edmund H. Smith,
Director of the Pacific Marine Station, Dillon Beach,
California for the use of the facilities and laboratories.
Page 52
To the staff of the Marine Station my thanks are due for
assistance in collecting.
This work was made possible by a grant from the De-
partment of the Interior (WP 010610), and by the finan-
cial help to the Coordenagéo do Aperfeigoamento de
Pessoal de Nivel Superior (CAPES), Rio de Janeiro,
Brasil.
LITERATURE CITED
AssoTtT, RoBERT TUCKER
1954. | American seashells. Princeton, New Jersey, D. van
Nostrand Co., Inc.; xiv+541 pp.; 100 text figs.; 40 plts.
ATKINS, DAPHNE
1937. On the ciliary mechanisms and interrelationships of
lamellibranchs. Part II. Sorting devices on the gills. Quart.
Journ. Micr. Sci. 79: 339 - 373
Part III. Types of lamellibranch gills and their food currents.
Quart. Journ. Micr. Sci. 79: 375 - 421
Boss, KENNETH JAY
1965. Symbiotic erycinacean bivalves. Malacologia 3 (2):
183 - 195
Deroux, G.
1961. Rapports taxonomiques d’un Leptonacé non décrit “Lep-
ton subtrigonum” JEFFREYS (nomen nudum — 1873). Cah.
Biol. Mar. 2 (2): 99 - 153
LeEsour, Marie V.
1938. The life history of Kellia suborbicularis.
Biol. Assoc. U. K. 22: 447 - 451
KEEN, A. Myra
1937. An abridged check list and bibliography of west North
American marine Mollusca. Stanford Univ. Press, Stanford,
Calif.; pp. 1 - 88; 2 figs. (29 September 1937)
MacGinitTie, Georce Esper & Nettie MacGInirtizE
1949. Natural history of marine animals.
McGraw-Hill, New York.
Narcui, WALTER
1966. The functional morphology of Ceratobornia cema, new
species of the Erycinacea (Mollusca, Eulamellibranchiata) .
An. da Acad. Brasileira de Ciéncias 38 (3/4) : 513 - 524
Journ. Mar.
473 pp.; illus.
THE VELIGER
Vol. 12; No. 1
OLpFIELD, E.
1955. Observations on the anatomy and mode of life of
Lasaea rubra (Montacu) and Turtonia minuta (Fasrictus).
Proc. Malacol. Soc. London 31 (5, 6): 226 - 249
1961. The functional morphology of Kellia suborbicularis
(Montacu), Montacuta ferruginosa (MonTAGU) and M. sub-
striata (MontTacu). Proc. Malacol. Soc. London 34 (5) :
255 - 295
1964. | The reproduction and development of some members of
the Erycinidae and Montacutidae (Mollusca, Eulamellibranchi-
ata). Proc. Malacol. Soc. London 36 (2): 79 --120
Oxproyp, IDA SHEPARD
1924. The marine shells of the west coast of North America.
Stanford Univ. Publ. Geol. Sci., 1: 1 - 247; plts. 1-57
OwEN, GARETH
1956. Observations on the stomach and digestive diverticula of
the lamellibranchia II. The Nuculidae. Quart. Journ. Micr.
Sci. 97 (4): 541 - 567 =n
PALMER, KATHERINE VAN WINKLE
1958. ‘Type specimens of marine mollusca described by P. P.
Carpenter from the West Coast (San Diego to British Colum-
bia). Memoir 76, Geol. Soc. Amer. i- viii + 1-376; plts.
1 - 35. New York, N. Y. (8 December 1958)
Poruam, M.L. ;
1940. The mantle cavity of some of the Erycinidae, Mont-
acutidae and Galeommatidae with special reference to the cili-
ary mechanisms. Journ. Mar. Biol. Assoc. U. K. 24: 549 - 587
Purcuon, R. DENISON
1958. The stomach in the Eulamellibranchia; Stomach Type
IV. Proc. Zool. Soc. London 131: 487 - 525
Quay te, D. B.
1960. The intertidal bivalves of British Columbia. Brit.
Col. Province. Mus., Dept. Educat. Handbook 17, 104 pp.
printed by Don MacDiarmid
Rerp, Rosert G. B.
1965. The structure and function of the stomach in bivalve
Journ. Zool. 147: 156 - 184
SrasEK, CHARLES ROBERT
1963. Synopsis and discussion of the association of ctenidia and
labial palps in the bivalved Mollusca. The Veliger 6 (2):
91-97; 5 text figs. (1 October 1963)
molluscs.
Vol. 12; No. 1
THE VELIGER
Page 53
Observations on Pervicacia tristis (DESHAYES, 1859)
and a Comparison with other Toxoglossan Gastropods
W. B. RUDMAN
Department of Zoology, University of Auckland
New Zealand
(5 Text figures)
INTRODUCTION
THE TOXOGLOSSAN PROSOBRANCH MOLLUSCS are a group
of specialised carnivores typified by a powerful neurotoxic
secretion and harpoon-shaped radular teeth. The most
studied family is the Conidae with work by Koun, CrEr-
NoHorsky and others. The cones studied so far are all
active selective predators, catching molluscs, worms and
fish (Koun, 1959, 1956, 1955 - 1956). This family is not
present in New Zealand, but the toxoglossans are repre-
sented by the Turrids, which are generally uncommon and
of small size, and the terebrid-like genus Pervicacia which
is also reported from Southern Australia and Japan. The
animal investigated is Pervicacia tristis (DESHAYES, 1859),
a member of the infauna of silty sand banks at low water.
Specimens were collected on the east coast, north of
Auckland at the Whangateau Harbour and at the mouth
of the Wade River. This animal lives just below the sur-
face of the sand, and often is found under small mounds.
Pervicacia lives in association with two tube worms,
Owenia fusiformis CutajE, 1844 and Axiothella quadri-
maculata AUGENER, 1914, and the molluscs Marginella
pygmaea Sowersy, 1846, Paratrophon stangeri (Gray,
1843), Zeacumantus spp., and Baryspira spp. Accompa-
nying these species is a small cumacean crustacean, prob-
ably of the genus Diastylis, which Pervicacia could pos-
sibly feed on. The animals collected refused to eat in
captivity and the gut contents were unrecognisable.
The animals were usually examined alive and dissected
under a binocular microscope. For histological and ana-
tomical investigation animals were fixed in Bouin’s fixa-
tive and the sections cut were stained with either Weigert,
van Giesen or Mallory Heidenhain stains.
EXTERNALS — SHELL anp BODY
Pervicacia tristis has a slender auger-shaped shell at-
taining 20 mm in height and 6 mm in width. Few animals,
however, reach this size. The coiling is dextral and radial
ribbing is often present, sometimes being very pronounced.
There is a horny operculum which fills the aperture when
the animal retracts. The body is translucent, with many
small white blotches showing under the surface formed
by a granular secretory substance found throughout the
foot and mantle regions.
The mantle cavity is wide and extremely deep, water
being drawn into it through the long inhalant siphon and
tested by the large osphradium before running over the
gills and passing out through an exhalant groove. The
large brown bipectinate osphradium lies on the extreme
left of the mantle cavity. To the right of this organ is
the long monopectinate ctenidium, each lamina being
triangular with one side being fixed to the mantle, ex-
tending to the back of the mantle cavity. The genital
duct opens also on the right, near the mantle edge in the
female, and into the penis in the male. To the left of
this organ is the rectum opening by a small anus. Between
the long black rectal gland and the ctenidium is the large
hypobranchial gland which produces a reddish purple
secretion. There are neither eyes nor tentacles. The sexes
are separate, the female duct ending in a large muscular
ring and the male duct in a long flat penis. The lack of
; val
Page 54 THE VELIGER Vol. 12; No
testis
kidney
digestive gland
vesicula seminalis oe
intestine
prostate gland
anus
penis
seminal duct
Vol. 12; No. 1
THE VELIGER
Page 55
eyes and tentacles, the large osphradium and the increase
in activity at night suggest that Pervicacia is nocturnal,
locating food by chemosensory means (Figure 1).
ALIMENTARY CANAL
On the ventral surface of the “head” or “snout” is an
opening, bordered longitudinally by a pair of muscular
lips, which forms the apparent mouth. This opening leads
into an anterior cavity which encloses the distal region of
the proboscis bearing the true or inner mouth. The “snout”
region, as is discussed later, is probably homologous with
the pseudoproboscis of the terebrids (Figures 1, 4 P). The
walls of the anterior cavity have a simple glandular epi-
thelium overlain by a thin layer of transverse (or radial)
muscle fibre. This is surrounded by a layer of muscle and
connective tissue. In Hastula cinerea (Born, 1780) an
“anterior tube” has been described, which when everted,
corresponds to this region (Marcus & Marcus, 1960).
In all specimens, whether alive or preserved, this region
was always extended and appeared incapable of with-
drawal. The buccal tube, or proboscis, which is only at-
tached to the body wall by a few small muscles, is wide
and highly muscular. Underlying the simple epithelium
lining the lumen, is a thin layer of longitudinal muscles,
followed by an ordered array of circular and transverse
(or radial) muscle blocks. On the floor of the buccal
tube is the odontophore, bearing a radula with two rows
of simple triangular teeth (formula 1:0°0-0:1). The
radular sac is short. Odontophoral cartilages are present,
contradicting FRETTER & GraHAm’s (1962) statement
that these have been lost in the Toxoglossa. The cartilag-
inous blocks are moved by two well-developed muscles,
M, and M2. When M, contracts and M: relaxes, the odon-
tophore rocks forward and when M, is relaxed and M2
contracts, the cartilage rocks backwards (Figure 4 S).
The apparently single salivary gland runs forward,
splitting in two. The ducts from each lobe join, forming
a common duct which enters the buccal tube near the
mouth (Figure 4 P). The oesophagus recurves sharply
twice, immediately behind the buccal tube, and then passes
through the nerve ring. The lining at the anterior end of
the oesophagus is ciliated and slightly folded, while
further back the folding increases and the layer of longi-
(<— on facing page)
Figure 1
Pervicacia tristis (DESHAYES, 1859)
Male, with mantle cavity opened
tudinal muscle is replaced by a larger one of circular
muscle. It is difficult to demarcate the transition from
oesophagus to stomach, but the stomach can be distin-
guished by heavier foldings of its wall. The gut wall here
contains odd groups of globules staining bright red with
Mallory Heidenhain and there is a thick layer of circular
muscle. Two large ducts leave the digestive gland, merging
to enter the stomach by a single opening. The histology
of the digestive gland is similar to that of some stenoglos-
sans described by FRETTER & GRAHAM (1962).
The intestine can be divided into two regions. Near the
stomach, red staining areas of granular cells are abundant
in the intestinal wall. The diameter of the lumen of the
intestine increases, almost all the folding disappears, and
the wall becomes very thin. This region of the intestine
is heavily ciliated and is the widest region of the alimen-
tary canal. The pallial or distal region of the intestine is
quite different. Near the back of the mantle cavity, the
intestine narrows and becomes triangular in cross-section,
the wall having large areas of red-staining granules. It has
no musculature and runs through the mantle as a narrow,
thick-walled, folded tube. It opens near the edge of the
mantle cavity and is followed down the mantle by the
rectal or anal gland (Figures 1, 3B).
REPRODUCTIVE SYSTEM
The ovary is a lobulated organ overlying the digestive
gland in the top whorls of the body. The ova stain grey
with Weigert, van Giesen stain. Part of the ovary is
packed with bright yellow staining oocytes. The genital
duct opens into the mantle cavity just behind the anus
and is modified into several specialized regions: albumen
gland; ingesting gland; capsule gland; and glandular
vestibule. The ovarian duct, leading from the ovary to the
albumen gland, is comparatively short, and is lined with
a simple epithelium. From it is a duct to the pericardial
cavity. The albumen gland has greatly folded walls, lined
with secretory and ciliated cells. Below the albumen
gland is the capsule gland (Figure 2 U). At the junction
of the two, a duct leads off to the “ingesting gland.” The
capsule gland is a large secretory region in which the duct
has become extended to form a narrow dorso-ventral slit
running forward in the mantle to open anteriorly into a
large distensible vestibule with folded muscular walls
(Figure 3B). This vestibule is a long muscular groove
opening into the mantle cavity. The muscular opening
of the groove could be considered the vagina. There is
no bursa copulatrix.
The length of the penis and the fact that it opens at
its tip suggests that this organ enters the capsule gland
Page 56 THE VELIGER Vol. 12; No. 1
testis
ves. sem.
prostate gland
nucleus ®)
iduct
i uc ®
albumen gland
cap. gl.
x st IK <p : 7 alae
a N\\ che as
seminal duct SO
eile
er blood sinus <
vest.
Vol. 12; No. 1
during copulation. The duct to the “ingesting gland” has
a glandular lining and is surrounded by a layer of circular
muscle. The cells of the epithelium are undifferentiated
and the duct often contains large numbers of oriented
sperm. This duct to the “ingesting gland” has been con-
sidered by FRETTER to be homologous with the receptacu-
lum seminalis of lower gastropods. It leads into a sausage-
shaped gland which in the live animal has a brown outer
covering. The gland is lined with vacuolated cells con-
taining groups of spherules staining brown with Mallory,
Heidenhain stain, vacuoles apparently being a site for the
ingestion of sperm. Unoriented sperm is found loose in
the lumen of the gland and also caught against the walls
of the gland cells. There is a layer of connective tissue sur-
rounding the gland cells. The duct and gland both have a
fairly thick layer of circular muscle. This gland is no doubt
the ingesting gland described by Fretrer (1941). The cap-
sule gland is a secretory region where the genital duct has
been extended into a narrow dorso-ventral slit. There are
two distinct glandular regions: a dorsal region and a
lateral one. At the ventral side of the lumen (Figure 3 B)
there is a loop to the animal’s left (i. e. a loop towards the
centre of the mantle cavity) which at its anterior end
opens into the vestibule. In FretrTer’s account of the
genital ducts of four prosobranchs (FRETTER, 1941) she
describes left and right longitudinal folds. Pervicacia and
Hastula (Marcus, 1960) have only a left fold. The lateral
glandular region is the most prominent area and consists
of groups of cells at different heights packed together in
lobes. The ducts of these secretory cells run parallel to
one another to open into the lumen of the gland through
the columnar ciliated epithelium. The dorsal lobe of
glandular tissue appears to be mucus-secreting and has a
much lighter appearance than the lateral lobe. The female
duct ends in a terminal pouch, similar to that described by
Marcus & Marcus (1960) in Hastula cinerea.
The reproductive system of the male can be divided
into six regions: testis; testis duct; vesicula seminalis;
prostate gland; vas deferens; and the penial duct.
(<— on facing page)
Figure 2
Pervicacia tristis (DESHAYES, 1859)
P — male reproductive system; Q — transverse section of prostate
gland; R — transverse section of vesicula seminalis; S — transverse
section of tip of penis; T — transverse section of penis; U — female
reproductive system: albumen gl. = albumen gland; b.s.s. = brown
staining spherules; cap gl. = capsule gland; g.p.d. = gonopericar-
dial duct; ig. = ingesting gland; prostate g. = prostate gland;
ves. sem. = vesicula seminalis; vest. = vestibule
THE VELIGER
Page 57
The testis is a large, greatly folded sac lying on the
digestive gland in the top whorls of the body. From this
organ runs a short duct, the testis duct, which soon widens
to form the much convoluted vesicula seminalis. This
runs down on the columella side of each whorl, finally
narrowing to a sphincter as it enters the pallial cavity.
The duct then flattens and is surrounded by glandular
cells to form the prostate gland running along the dorsal
side of the body. The duct then narrows and becomes sur-
rounded with a layer of circular muscle. This is the true
vas deferens and runs forward until it enters the penis
just to the right of the snout. The duct passes down the
penis and opens at the tip (Figure 2 P). The structure of
the vesicula seminalis is the same as that of Ocenebra
(FretTer, 1941). The epithelium consists of columnar
cells which may be ciliated, but when the organ is filled
with sperm, such an observation is impossible. The nuclei
of the cells are large, being either spherical or irregular
in shape, and the cytoplasm is vacuolated and contains
large yellow-brown staining spherules similar to those in
the ingesting gland. It is thought that the vesicula semi-
nalis also ingests sperm either to keep the duct clear or to
remove old sterile sperm (Figure 2R). The character
of the epithelium alters when the vesicula seminalis enters
the pallial cavity, becoming quite simple. The vesicula
seminalis leads to the prostate gland situated on the right
dorsal side of the body. These two regions are separated
by a sphincter.
The prostate gland is a flattened region of the duct,
the lateral walls being greatly thickened by the growths
of glandular cells. These glandular regions on each side
of the longitudinal ciliated duct differ from each other
in many ways. The most obvious distinction is the colour
the regions stain. The left or upper glandular region stains
red, suggesting a protein or nutritive secretion, while the
right dorsal region stains a light blue with Mallory Hei-
denhain, suggesting a mucous secretion. On the ventral
side of the duct a glandular region of large columnar
cells is present. The function of these cells is unknown.
The lumen is ciliated on its ventral side. The vas deferens
is a small heavily ciliated tube surrounded by a thick
coat of circular muscle. This duct runs into the penis and
travels forward on one side of the penis until it reaches
the penial opening (Figure 2 Q).
The penis is a long muscular organ half the length of
the animal’s long axis (Figure 1). It lies folded within
the mantle cavity, when not in use, and has an opening
at its terminal end through which sperm can be ejected
during copulation. The glandular epithelium and mus-
culature of the penis (Figure 2 T) is extremely complex.
There is an outer covering of secreting cells probably to
ease passage at copulation. Inside this is a layer of ob-
Page 58 THE VELIGER Vol. 12; No. 1
Si
SS
rectal
gland
capsule %
gland <£:.
osphrad.
0.0. 5 64.0
"+. columellar muscle . 7... |
vestibule Bao Waae SOP ons
09 5,00
Vol. 12; No. 1
lique muscle surrounding a very conspicuous layer of
transverse and longitudinal fibres. The centre of the penis
is loosely packed with connective tissue penetrated by
blood spaces. One side of the penis carries the penial
seminal duct, on the other there is an open space which
may be a blood channel. The penial nerve, a derivation of
the pedal nerve, is situated off-centre. Proximally the semi-
nal duct opens into a wide ciliated groove which becomes
W-shaped (Figure 2S). Both grooves and the central
ridge are highly ciliated. This groove is probably a
vestige of a primitive open groove along the penis. The
blocks of transverse and longitudinal muscle in the penis
suggest that peristalsis effects the ejection of sperm.
COMPARISON
A number of terebrids collected by Professor J. E. Mor-
ton during the Royal Society Expedition to the Solomon
Islands, 1966 were kindly given to the author and a study
of their gross anatomy was undertaken, The results of
this are listed below:
Terebra babylonia. This animal possesses eyes on very
short stalks and its reproductive system, in both sexes, is
similar to that of Pervicacia tristis. The foregut, however,
is quite different. A muscular cylinder extends back into
the anterior cavity opening at its inner end through a
slit bordered by a pair of muscular lips. This slit is no
doubt “la fente dorsale” of Bouvier (1887) and RisBEc
(1953). The muscular cylinder has been called by RisBEc
(op. cit.) “la gaine de la trompe” — the proboscis sheath —
in his discussion on Terebra muscaria LAMaRCK, 1822, and
by Bouvirr, “la gaine proboscidienne.” This structure,
however, is quite different from the proboscis sheath of
the lower stenoglossans and a new term should be em-
ployed. Marcus & Marcus (1960) have studied the te-
rebrid Hastula cinerea and use the term “anterior tube.”
This term, I feel, is unsatisfactory and suggest instead
“pseudoproboscis.” Behind the pseudoproboscis, and quite
separate from it, is the true or inner mouth, the anterior
opening of the alimentary canal. The proboscis is long
and at its inner end passes through a thin wall separating
the anterior cavity from the true body cavity. Inside the
body cavity proper, a small radular sac opens into the pro-
(<— on facing page)
Figure 3
Pervicacia tristis (DESHAYES, 1859)
A — copulation; B — transverse section through mantle cavity:
hyp. gl. = hypobranchial gland; osphrad. = osphradium; o.n. =
osphradial nerve; oes. = oesophagus; sal. g. = salivary gland
THE VELIGER
Page 59
boscis. Also opening at this point are the salivary gland
ducts and the poison gland. The long thin poison gland
folds around the oesophagus and the posterior end of the
proboscis, leading to the large muscular bulb often mistak-
enly identified as the poison gland. The oesophagus, on
leaving the proboscis, folds forward under the anterior
cavity, before folding back to lead posteriorly to the
stomach. The radular teeth are harpoon-shaped, the
chitin being formed into a rolled, hollow tube (Figure
Ai@ wily)
Terebra maculata and Terebra affinis. These animals were
also collected at the Solomon Islands and as with the above
species, possess eyes on small stalks and have a reproductive
system similar to that of Pervicacia tristis. However, the
foregut again differs considerably in structure. In both
these species the pseudoproboscis is extremely long (in
T. maculata it lies folded in the anterior cavity), while
the true proboscis is a small remnant devoid of radula.
There is no poison gland and the salivary glands are much
reduced. The oesophagus leads from the proboscis, folding
under the pseudoproboscis before dropping through the
ventral wall of the anterior cavity into the true body
cavity. Terebra dimidiata was also found to have a very
similar structure (Figure 4 R).
For the purposes of discussion short accounts of other
terebrids already studied are listed below.
Hastula cinerea. The reproductive system is similar to
that of Pervicacia tristis except for the possession of a
small bursa copulatrix near the capsule gland. The fore-
gut is similar to that of T: babylonia, the eversible pseudo-
proboscis, however, being shorter in relation to the pro-
boscis (Marcus & Marcus, 1960).
Terebra muscaria. There is neither radula nor poison
gland (Rispec, 1953).
Terebra cancellata. This species has both poison gland
and harpoon-shaped radular teeth (RisBec, 1953).
DISCUSSION
The foregut of the Terebridae is fundamentally different
from that of the Conidae and some of the Turridae, in
which the alimentary canal opens externally, there being
no anterior cavity. In Cenodagreutes aethus E. H. SmitH,
1967, and C. coccyginus E. H. Situ, 1967, two turrids
recently described from England (E. H. Smirn, 1967)
there is neither radular nor poison gland. In these animals
the oesophagus is continuous with the large proboscis
forming the rhynchodaeum, and opens directly to the out-
side. In Mangelia brachystoma (Pui.ippi, 1844) a turrid
studied by Ropinson (1960) the proboscis, as in the
terebrids, opens into an anterior cavity, which in turn
Vol. 12; No. 1
THE VELIGER
Page 60
anterior cavity
proboscis
pseudoproboscis
i=)
fo}
Py
sac
‘.-—— oesophagus
proboscis
pseudoproboscis
cartilage
“ett
Ra
=
xeN
Vol. 12; No. 1
opens to the exterior. Mangelia differs from the terebrids
in not having a pseudoproboscis. The anatomy of the
Conidae is not well-known. The reproductive system of
Conus mediterraneus BruGuIERE, 1792 has been studied
(Martoja-Prerson, 1958) while Koxn’s studies on
feeding and the foregut of this family suggest that the
foregut of the cones has more in common with the Tur-
ridae than with the Terebridae.
It is considered that the Recent families of the Toxo-
glossa: Terebridae, Conidae, and Turridae, are a closely
related group of advanced gastropods (Powe Lt, 1966).
However, there is little known of their anatomy. PowEtt,
whose comprehensive reclassification of the Turridae, on
shell and radular characters, forms a massive contribution
to our knowledge, and Kou studying in detail the feeding
and ecology of the Conidae are almost the only contem-
porary students of the Toxoglossa. Shorter papers of vary-
inf usefulness, have dealt with aspects of the anatomy
and functional morphology of a few members of the
group. From what is known of the toxoglossan reproduc-
tive system, Pervicacia tristis differs only slightly from the
general pattern. In terebrids, the pallial vas deferens leads
into a swollen prostate gland and from this to the tip of a
long penis. As Marcus (1960) has shown in Hastula
cinerea, in Pervicacia tristis, Terebra affinis and T. macu-
lata, this duct is closed (personal observations). RisBEc
(1953) states that in Terebra muscaria there is an open
duct. In two turrids examined, Mangelia brachystoma by
Roginson (1960) and Phenatoma zealandica(E. A. SmirH,
1877) by Ponder (personal communication) there is no
swollen prostate region. Although no bursa copulatrix
exists in P. tristis, a small sac near the capsule gland in the
turrids Hastula cinerea and Cenadogreutes spp. (E. H.
SmirH, 1967), and a large sac at the genital opening of
Conus mediterraneus studied by Martoja-Pierson, 1958,
suggest that this organ is present in varying degrees
throughout the Toxoglossa.
This study has shown that the foregut of the members
of the Terebridae has developed a peculiar structure, the
pseudoproboscis. In Hastula cinerea this structure is small
and is everted during feeding (E. « E. Marcus, 1960).
(<— on facing page)
Figure 4
Pervicacia tristis (DESHAYES, 1859)
P — sagittal section through head region U -— dorsal view of
odontophore; V — tooth; S — section through odontophore.
Terebra babylonia: Q - sagittal section through head region;
T — tooth.
Terebra affinis: R — sagittal section through head region.
THE VELIGER
Page 61
When this tube is not everted (that is, lying in the anterior
cavity), the opening at its inner end is near the opening
of the alimentary canal. In Terebra maculata this opening
of the pseudoproboscis is a ventral slit bordered by a pair
of muscular lips, and is similar in structure to the opening
of the anterior cavity of Pervicacia tristis. In T. babylonia
the opening is in a central position while in H. cinerea it
is dorsally placed. The size of the pseudoproboscis differs
from species to species; in Hastula the tube is short, the
true proboscis long and folded and there are both poison
gland and harpoon-shaped radular teeth. Terebra baby-
lonia is similar although the pseudoproboscis is relatively
longer. In the three species studied having neither radular
nor poison gland the pseudoproboscis is very long and in
T. maculata the length is so great that the tube is folded
about itself when lying in the anterior cavity. In contrast
to T. babylonia the true proboscis in these forms is ex-
tremely small, being merely a short muscular vestibule at
the end of the oesophagus. There is little doubt that in
these cases the prey is caught, not by the “typical” toxo-
glossan method of harpooning the prey with a poison
tooth, but by the long pseudoproboscis being everted and
sucking in the prey (Figure 5 G). One difficulty in under-
standing this rather unique method of food collecting is
the question of how food is transported from the pseudo-
proboscis to the opening of the true proboscis with which
there is no direct connection. This problem cannot be
solved with preserved material and must wait a study
of living animals.
The one character which distinguishes the Toxoglossa
from all other gastropods is the possession, by some mem-
bers of each family, of a poison gland and harpoon-shaped
radula teeth. As Powe. (1966) has suggested, it is most
probable that the Conidae and the Terebridae are special-
ised offshoots from turrid stock. The Turridae possess a
wide range of radular teeth with formulae ranging from
the prototypic 1-1-1-1-1 of Drillia and Clavus, where
the laterals are wide and serrate, to the advanced 1-0-0
‘0-1 harpoon-shape of Phenatoma (Figures 5A-D). In
all cases studied by Ponper, a poison gland was present,
even in the cases where a prototypic radula was present.
Only in the most advanced forms was a cartilaginous
odontophore absent (W.F Ponder, personal communi-
cation). The genus Cenodagreutes is unique in the turrids,
the radula, poison gland and salivary glands being com-
pletely absent (E.H.Smiru, 1967). The generic name
meaning “the toothless hunter” is very apt. It is difficult
to agree with PoweELt’s proposed subfamily classification
of the Turridae. In three of the subfamilies, Clavinae,
Mangeliinae, and Clavatulinae, there are species possessing
the advanced harpoon-shaped radular teeth and species
possessing more primitive radular forms. Considering
Page 62 THE VELIGER Vol. 12; No. 1
TURRIDAE
TEREBRIDAE
CONIDAE
PERVICACIIDAE :
(h) (b)
@ SPEIGHTIIDAE
Vol. 12; No. 1
that subfamilies should represent common lines of des-
cent, if we are to agree to PowELL’s groupings, then we
must also agree that harpoon-shaped radular teeth have
arisen independently on three different occasions. Because
of the close relationships of this group this does not seem
reasonable.
In the Conidae all species that have been studied have
a specialised foregut with a poison gland and an advanced
radular tooth (Figure 5 L). With the exception of Per-
vicacia tristis, the terebrids fall into two categories. One
group has a foregut similar to that of the cones, while
the other has neither poison gland nor radula. It is sug-
gested that the latter group has lost these organs. It is
therefore possible to suppose that, with the exception of
Pervicacia, both the Conidae and the Terebridae arose
independently from turrids that had developed harpoon-
shaped radular teeth.
The genus Pervicacia is unique within the Toxoglossa,
having no poison gland and having a primitive type of ra-
dula. Although the shell is terebrid in shape, no valid phylo-
genetic argument could continue to place it in the Tereb-
ridae. A new family within the Toxoglossa is therefore
proposed. :
Pervicaciidae new family.
Toxoglossan gastropods bearing a terebrid-like shell,
axially ribbed. The shell is auger-shaped, less highly calci-
(<— on facing page)
Figure 5
Suggested phylogenetic relationships within the Toxoglossa
Turridae:
A — Drillia umbilicata (Gray, 1838), radula
B — Rhodopetoma rhodope (DALL, 1919), radula
C - Haedropleura septangularis (MoNnTAGU, 1803), radula
D — Phenatoma zealandica (SmitH, 1877), radula
E —- Phenatoma zealandica (Smiru, 1877), animal
Terebridae:
F - Hastula cinerea (Born, 1780), radula
G - Terebra dimidiata (LINNAEUS, 1758), ingesting worm
Pervicaciidae:
H — Pervicacia tristis (DESHAYES, 1859), radula
I — Pervicacia tristis (DESHAYES, 1859), animal
Speightiidae:
J — Speightia spinosa (SuTER, 1917)
Conidae:
K — Conus striatus LINNAEUS, 1758, ingesting fish
L — Conus arenatus BRuGUIERE, 1792, radula
(A, B, C, after PowELt, 1964; F, after Marcus, 1960; K, partly
from photo, Koun, 1956; L, after THIELE, 1931)
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Page 63
fied than in the Terebridae. Radula primitive, 1:0:0-0-1,
odontophoral cartilage blocks present. Poison gland absent.
Type genus: Pervicacia IREDALE, 1924.
It is doubtful if the family Pervicaciidae is even toxo-
glossan, but the similarity of the reproductive system, and
because the radula is similar to that of some primitive
turrids, it was decided to leave it within the superfamily
Toxoglossa.
It is suggested that sometime before the Cretaceous a
gastropod developed a single poison gland and a radula
perhaps similar to that of Drillia umbilicata (Gray, 1838),
and through gradual loss of the central and marginal
teeth the turrids developed the highly specialised radular
form of Phenatoma zealandica (E.A.SmituH, 1877).
Early in this development a form such as Pervicacia could
have split off leading, with loss of the poison gland, to the
genus Pervicacia. Both the Conidae and the Terebridae
split from the turrid line after the development of the
harpoon-shaped radular teeth. Some terebrids have speci-
alised further than the cones, members having lost both
the poison gland and the radula. One fossil family placed
with the Toxoglossa, the Speightiidae, disappears from
the fossil record at the end of the Eocene (PowELt, 1966)
and their inclusion within the superfamily is purely con-
jectural. As mentioned earlier, Powe w’s internal classi-
fication of the Turridae must remain open to doubt
until more than the shell and radula is known in this
family (Figure 5).
The similarity of shell between the Terebridae and the
Pervicaciidae is not unparalleled in the Toxoglossa, the
Conidae having a similar shell-shape to that of Conorbis
dormitor SOLANDER, 1766, a supposed turrid. Within the
cones certain plasticity is seen in the shell, the fast moving
piscivorous forms, such as Conus geographus LINNAEUS,
1758, have lightly calcified, wide apertured shells while
the slower vermivores have heavy shells with slit-like
apertures. It is reasonable to assume that, through living
in similar habitats, the terebrids and the pervicacids have
developed similarly shaped stream-lined shells.
SUMMARY
A study of Pervicacia tristis (DESHAYES, 1859) with
special reference to the gut and reproductive system.
The possession of a cartilaginous odontophore and prim-
itive radula is reported,The foregut region of a number
of tropical terebrids: Terebra maculata Linnaeus, 1758,
T. dimidiata Linnaeus, 1758, T: babylonia LaMarck,
1822, and T. affinis Gray, 1834 is described. These species
are compared with some other toxoglossans whose ana-
Page 64
tomy has been studied previously. The genus Pervicacia is
removed from the Terebridae and a new family is proposed
to contain it.
ACKNOWLEDGMENTS
I would like to thank Professor J. E. Morton, Head of
the Zoology Department, for making available some of
his collected material for study; to Dr. M. C. Miller for
his encouragement, and to the Royal Society, London,
who sponsored the Solomon Islands Expedition on which
the terebrid material was collected.
LITERATURE CITED
ALLAN, Joyce
1959. Australian shells; rev. ed.
487; plts. 1-44; text figs.
Bouvier, E. L.
1887. Systeme nerveux, morphologie générale et classification
des gastéropodes prosobranches. Ann. Sci. Nat. Zool., ser.
7, vol. 3 Paris.
CERNOHORSKY, WALTER OLIVER
1964. The Conidae of Fiji (Mollusca: Gastropoda). The
Veliger 7 (2): 61-94; plts. 12-18; 3 text figs.; 1 map
(1 October 1964)
Melbourne, i- xxi; 1 to
FRETTER, VERA
1941. The genital ducts of some British stenoglossan proso-
branchs. Journ. mar. biol. Assoc. U. K. 25: 173 - 211
FRETTER, VERA, & ALASTAIR GRAHAM
1962. British prosobranch molluscs, their functional anatomy
and ecology. London, Ray Soc. xvi + 755 pp.; 316 figs.
Jouansson, J.
1953. On the genital organs of some mesogastropods. Zool.
Bidr. Uppsala, 30: 1 - 23
THE VELIGER
Vol. 12; No. 1
Koun, ALan Jacoss
1955. Studies on food and feeding of the cone shells. Rep.
Bull. 22, Amer. Malacol. Union
1956. Feeding in Conus striatus and C. catus. Proc. Hawai.
Acad. Sci. 31st. ann. meetg., 1955-56. Univ. Hawaii publ.
1956. — Piscivorous gastropods of the genus Conus. Proc.
Nat. Acad. Sci. 42 (3): 168-171
1963. | Venomous marine snails of the genus Conus; in: Ven-
omous and poisonous animals and noxious plants of the Pa-
cific area. Pergamon Press, Oxfr., 83-96; 4 text figs.; 3 tab.
Lesour, Marte V.
1937. The eggs and larvae of the British prosobranchs with
special reference to those living in the plankton. Journ.
Mar. Biol. Assoc. U. K. 22
Marcus, EvELINnE pu Bors-REYMoND & ERNST Marcus
1960. On Hastula cinerea. Bol. Fac. Fil. Gen. Letr. Univ.
S. Paulo, Brasil, no. 260. Zoologia no, 23: 25 - 66
Martoya-Prerson, M.
1958. Anatomie et histologie de l'appareil génital de Conus
mediterraneus Brug. Bull. Biol., fasc. 2: 1 - 22
Oswa Lp, A.
1893. Der Riisselapparat der Prosobranchier.
schr. Natur. 28: 119 - 162
PowELL, ARTHUR WILLIAM BADEN
1964. Family Turridae in the Indo-Pacific. Part 1.
Pacific Moll. 1, no. 5
RIsBEc, JEAN
1953. | Observations sur l’anatomie des Terebridae néocalédoni-
ens. Bull. Mus. Nat. Hist. nat. Paris 25: 576 - 583
RosBINSON, ELIZABETH
1960. Observations on the toxoglossan gastropod Mangelia
brachystoma (PHILiPP!). Proc. Zool. Soc. London 135:
319 - 338
SmitH, EpMuND Hosart
1967. | Two new species of British turrids.
10 (1): 1-4; plt. 1; 5 text figs.
THIELE, JOHANNES
1931-1935. Handbuch der systematischen Weichtierkunde.
Jena, pp. 1- 1154; 893 text figs.
Jen. Zeit-
Indo-
The Veliger
(1 July 1967)
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THE VELIGER
Page 65
A Possible ““‘Defense’”” Response in a Commensal Polychaete
BY
RONALD V. DIMOCK, Jr.'
AND
JOYCE G. DIMOCK
Department of Biological Sciences, The University of California, Santa Barbara, California 93106
(Plate 4)
INTRODUCTION
THE KEYHOLE LIMPET Diodora aspera (ESCHSCHOLTZ,
1833) is one of a diverse group of hosts (PETTIBONE,
1953) for the commensal polynoid Arctonoe vittata
(Gruse, 1855). This limpet occurs on rocks in the low
intertidal.zone of open coasts from Alaska to Lower Cali-
fornia (RicKETTS & Catvin, 1962). Limpets, 50 - 60 mm
long, may have commensal A. vittata as long as 50 - 70 mm
inhabiting the mantle cavity. PALMER (1968) has reported
a frequency of infestation of 0.8 to 1.0 worms per host
limpet.
Marco.in (1964) reported a distinct mantle response
from Diodora aspera which he suggested is a possible
deterrent against predation by sea stars. This escape res-
ponse thus might help protect both the limpet and its
commensal from predation. In this paper we report a
biting response by commensal Arctonoe vittata toward
potential asteroid predators which might in turn afford
an added measure of protection to the symbionts.
MATERIALS anp METHODS
Diodora aspera with commensals were collected during a
series of minus tides during August and September, 1968,
from the Edward’s Reef area on the southwest side of
San Juan Island, Washington. The common intertidal
asteroid, Pisaster ochraceus (BRANDT, 1835), was used as
the principal test animal for the determination of the Arc-
tonoe response. In addition, the reaction of commensal A.
vittata to Evasterias troschelii (Stimpson, 1862) and
* Supported under ONR Contract No. 4-222(03), D. Davenport,
Principal Investigator.
Pycnopodia helianthoides (BRANDT, 1835), two typically
subtidal sea stars, was tested. All animals were maintained
in aquaria with running sea water at the Friday Harbor
Laboratories.
The procedure for observing the biting response con-
sisted of placing a single Diodora containing a commensal
in an enameled tray of fresh sea water. The limpet was
left undisturbed in the tray until it had relaxed and was
moving about on its foot. An arm of a test star was then
brought into contact with the limpet in such a way that
the extended tubefeet contacted the lateral or posterior
shell-mantle junction. Keeping the rest of the sea star
free of the test site allowed observation of both the sym-
bionts. The trial period for these experiments was 10
minutes.
The behavior of isolated Arctonoe vittata toward Pis-
aster ochraceus was examined by merely placing single
worms in contact with a sea star in a tray of sea water.
To determine whether the mantle response of Diodora
results in the release of some substance which affects the
behavior of Arctonoe vittata toward Pisaster, individual
Arctonoe were removed from Diodora and placed next to
a Pisaster at one end of a 12-inch plastic tray. A Diodora
was then placed at the other end of the tray and the
mantle response elicited by a sea star. Sea water was
allowed to flow over the Diodora toward an outlet at
the end of the tray having the Arctonoe and Pisaster.
The necessity for the sea star’s presence in eliciting the
response of the commensal Arctonoe was examined by
subjecting individual limpets with their commensals to
sea water in which sea stars had been maintained prior to
testing. In these experiments Pisaster and Pycnopodia,
large enough to displace 500 ml, were placed in individ-
ual beakers of 4000 ml capacity, with aerated sea water
for 27 hours. The Diodora were placed in individual elass
Page 66
dishes (150 ml capacity) of fresh sea water for 5 minutes
prior to testing. For each test the water in the dish was
replaced with “sea star water” and the activity of the
limpets and commensals recorded during the 10 minute
trial period.
The feeding experiments were conducted in a 22 inch
by 82 inch Lucite tank 22 inches deep and divided into
two equal compartments. Ten Diodora containing Arc-
tonoe vittata were placed in one compartment (I) and
10 lacking Arctonoe in the other (II). In the first experi-
ment 3 Pisaster ochraceus (starved 3 days) of similar size
were placed in each compartment. In the second experi-
ment 2 Pycnopodia helianthoides of similar size were
placed in each compartment. Natural lighting conditions
illuminated the outdoor tank.
RESULTS
The Biting Response
Contact between Pisaster ochraceus tubefeet and the
Diodora aspera mantle edge immediately evoked the
“mantle response” described by Marcouin (1964). In
several preparations, activity of Arctonoe vittata was ob-
served within 15 seconds after initiation of the mantle
response. The first evidence of the activity of Arctonoe
usually consisted of observing the dark tipped palps of
the worm extending slightly past the mantle margin. This
initial behavior occurred whether or not the worm’s head
was in the immediate vicinity of the tubefoot contact.
Next, the worm usually would extend itself about 1 cm
from the edge of the host shell (Plate 4, Figure 1). If
the worm emerged away from the point of sea star
contact, it “searched” the immediate area with probing
movements. Such a worm would usually move around
the perimeter of the shell toward the point of tubefoot
contact. This movement could be accomplished either by
moving while extended or by retracting under the mantle
and emerging in the vicinity of the sea star’s contact.
In the vicinity of the tubefeet, the worm intensified its
searching behavior until actual contact was effected with
a tubefoot. At this point the worm typically explored the
tubefoot by moving its head up and down the length of
the foot with a palp on each side. Following this explora-
tion the worm rapidly everted its armed proboscis and
bit the tubefoot (Plate 4, Figure 2).
Several modifications in this behavior occurred. Occa-
sionally the worm would emerge from under the shell up-
side down. A number of worms were observed to bite the
sea star deep in the ambulacral groove rather than simply
THE VELIGER
Vol. 12; No. 1
on an exposed tubefoot. In one instance an Arctonoe vit-
tata bit an Evasterias troschelu on the lateral margin of
one ray well above the ambulacral groove.
The response usually was completed within 2 - 3 min-
utes following contact between Pisaster tubefeet and the
Diodora. Single bites occurred most frequently, but oc-
casionally repeated biting occurred during the 10 minute
experimental period.
The usual result of the biting behavior was the imme-
diate withdrawal of the tubefeet in the vicinity of the
worm, often including any tubefeet attached to the host’s
shell. A bite on the side of a ray frequently resulted in
the sea star’s bending the ray away from the point of
biting. At no time during these experiments did we see
a tubefoot severed nor did any worm exhibit a response
to an excised tubefoot held in forceps or a torn tubefoot
on the bottom of the tray.
Pisaster ochraceus consistently elicited more biting re-
sponses from Arctonoe vittata than did either Evasterias
troschelu or Pycnopodia helianthoides. In a series of 10
trials with each species, Arctonoe bit Pisaster 8 times while
biting both Evasterias and Pycnopodia 2 times each.
The Behavior of Isolated Arctonoe vittata toward Pisaster
ochraceus
When placed in contact with the tubefeet of Pisaster
ochraceus, Arctonoe removed from host limpets wandered
among the tubefeet and occasionally wrapped themselves
around the tip of a ray. In a series of 10 such trials the
only adverse interaction observed between the two species
consisted of the sea star’s pedicellariae impeding the at-
tempts of Arctonoe to climb onto a ray. At no time did an
isolated Arctonoe bite a sea star.
There was no observable change in the behavior of
isolated Arctonoe toward Pisaster when the commensals
and sea stars were placed together in the effluent from
a Diodora which was being harassed by a second Pisaster.
Though this experiment was repeated only 3 times, it
suggests that a water transportable substance is not re-
leased from Diodora during the mantle response which
might modify Arctonoe’s behavior toward Pisaster.
The Response to “Sea Star Water”
In a series of 10 trials in which water from Pycnopodia
was used, no mantle response of the limpets was observed
nor did any commensal Arctonoe become active. The
addition of water in which Pisaster had been maintained
elicited a very slight mantle response from Diodora in 9
trials and a pronounced response, approximately one-
half the response to direct tubefoot contact, in one limpet.
THE VELIGER, Vol. 12, No. 1 [Dimocx « Dimock] Plate 4
Figure 1
Arctonoe vittata extended from the mantle cavity of Diodora aspera
and approaching the tubefeet of Pisaster ochraceus.
Figure 2
The rapid eversion of its armed proboscis by Arctonoe vittata
immediately prior to biting a tubefoot of Pisaster ochraceus.
Vol. 12; No. 1
None of these 10 trials elicited any response from the
Arctonoe. Sea water controls had no noticeable effect on
either the limpets or their commensals.
Feeding Experiments
During the course of the Pisaster — Diodora feeding
experiment, Arctonoe were occasionally observed biting
Pisaster. These responses invariably resulted in a retrac-
tion of the tubefeet by Pisaster and frequently caused
the star to avoid the limpet it was in contact with at the
time of the biting. On the 10" day of the experiment, 2
minced Diodora were introduced into each compartment
in an attempt to stimulate feeding. The minced Diodora
were ingested, but upon termination of the experiment
at day 12, no Diodora had been eaten in either compart-
ment.
The experiment with Pycnopodia resulted in 7 (out
of 10) Diodora with Arctonoe (compartment I) and 5
(out of 10) without Arctonoe (compartment II) being
eaten within the week. At the end of this experiment (7
days) no free Arctonoe were observed in compartment I
although each of the 3 remaining Diodora had only one
Arctonoe with it. This suggests that 7 commensals were
ingested with their hosts. Indeed, examination of a Dio-
dora which had been ingested by a Pycnopodia in com-
partment I revealed the commensal still with its host.
DISCUSSION
The biting response of commensal Arctonoe does not
appear to be a simple feeding reaction elicited by the
presence of a sea star. The failure of any Arctonoe to
remove a tubefoot from a star or to respond to excised
tubefeet supports this observation. Similarly, the occasion-
al biting of an area distant from the tubefeet suggests a
need for another explanation.
The mechanisms involved in initiating the biting res-
ponse are not as yet clear. The marked difference in the
behavior of isolated worms toward Pisaster suggests that
host proximity is involved in the “attack” of the star by
Arctonoe vittata. The possibility of the existence of some
substance released during the mantle response which may
trigger this behavior in Arctonoe must be further investi-
gated. However, the physical changes in the worm’s en-
vironment associated with the mantle response may be
sufficient stimulus to initiate the commensal’s activity.
Our observations support Marcouin’s (1964) sugges-
tion that the mantle response of Diodora aspera is suffi-
cient to protect the limpet from Pisaster ochraceus. How-
ever, there is no evidence that Pisaster is normally a
predator on Diodora. Fever (1959) did not include D.
THE VELIGER
Page 67
aspera in the tabulation of organisms eaten in the field by
this sea star in the Monterey Bay area of California.
Marcouin (1964) did report, however, that Pisaster ate
Diodora when the mantle response was inhibited by an-
aesthesia.
Pycnopodia had no difficulty capturing and eating Dio-
dora aspera, suggesting that the mantle response is ineffec-
tive against predation by this agile, multirayed sea star.
Apparently, the presence of Arctonoe vittata had no effect
on the attractiveness of Diodora as a food organism. To
ascertain whether the presence of A. vittata has any
survival value for Diodora against predation by either
Pisaster or Pycnopodia, further experiments must be
conducted. A more pertinent experiment than those de-
scribed herein might entail presenting the stars with a
choice between Diodora with and without commensals.
These experiments could be coupled with experiments
offering alternate food species to the sea star. The tendency
of Arctonoe to move from host to host under laboratory
holding conditions would complicate such experiments.
Detailed observations of related symbioses may reveal
additional complex behavioral interactions suggestive of
mutualistic associations. Such interactions between sym-
bionts, mediated by more complex mechanisms than semi-
Passive one-way commensal interactions, might help ex-
plain the observed specificity (DAvENPoRT, 1966) and
the high infestations (PALMER, 1968) of various symbiotic
associations.
ACKNOWLEDGMENTS
The authors wish to express their appreciation to Dr.
Robert L. Fernald for providing research space at the
Friday Harbor Laboratories and to Mr. Richard Delingher
of the Museum of Comparative Zoology, Harvard Uni-
versity, who made the first observation at the Bodega
Marine Laboratory, University of California, and brought
the response to our attention. We especially wish to thank
Dr. A. O. Dennis Willows for assistance in making a short
film record of this response.
LITERATURE CITED
DAVENPORT, DEMOREST
1966. The experimental analysis of behavior in symbioses.
pp. 381 - 429, In: S.M. Henry, Symbiosis, vel. 1. Acad. Press,
New York
FepER, Howarp MITCHELL
1959. The food of the starfish, Pisaster ochraceus, along the
California coast. Ecology 40 (4): 721-724; 2 text figs.
(October 1959)
Page 68 THE VELIGER Vol. 12; No. 1
Marco.Lin, ABRAHAM STANLEY
1964. The mantle response of Diodora aspera. Animal
Behavior 12: 187 - 194
PaLMER, JoHN B.
1968. An analysis of the distribution of a commensal polynoid
on its hosts. Ph. D. thesis, Univ. Oregon; 121 pp.
PETTIBONE, Marian H.
1953. | Some scale-bearing polychaetes of Puget Sound and ad-
jacent waters. Univ. Washington Press, Seattle, Wash.,
136 pp.; illustr.
Ricketts, Epwarp F. & Jack CALVIN
1962. Between Pacific tides. 3rd ed., rev. by Jor W.
HEpGPETH. xiii+502 pp.; illustr. Stanford Univ. Press, Stan-
ford, Calif.
Vol. 12; No. 1
THE VELIGER
Page 69
Occurrence of the Cephalaspid Philine sinuata (Stimpson)
in Southern New England, with a Discussion of the Species
BY
D. R. FRANZ anno K. CLARK
Systematic and Environmental Biology, Biological Sciences, University of Connecticut, Storrs, Connecticut 06268
and
Marine Research Laboratory, University of Connecticut, Noank, Connecticut 06340
(8 Text figures)
Philine sinuata 1s A MINUTE tectibranch species described
by Writu1AM Stimpson (1850) on several specimens
dredged in shallow water from Boston Harbor, Massa-
chusetts. Although not figured in the original description,
a drawing of the shell was included in a subsequent pub-
lication (Stimpson, 1851) and this figure was copied by
Goup & Binney (1870). Although this species may be
fairly common in New England, there has been no further
published information on its biology or occurrence in
North America. The purpose of this brief report is to
note the occurrence of P sinuata in southern New England
and to present further information regarding its morphol-
ogy and taxonomy. These observations are based on ani-
mals collected during the summer of 1968 from a location
near the mouth of the Mystic River at Noank, Connecti-
cut (Fishers Island Sound). Further observations and
measurements were made on several lots of specimens
collected by William Clapp from the area of Duxbury,
Massachusetts. In this regard, the authors wish to ac-
knowledge the generosity of Dr. Joseph Rosewater who
provided us with the available collections on deposit at
the United States National Museum.
ECOLOGY
Like all philines, Philine sinuata is an infaunal species. The
Connecticut animals were collected from a shallow sub-
littoral station at a depth of about 3 m and a salinity of
‘ Contribution No. 53 of the Marine Research Laboratory, Uni-
versity of Connecticut.
from 29 - 30%.. Three of the four animals were picked off
the surface of the sediment by one of us using SCUBA
gear. The fourth specimen was taken with a Peterson
Grab. The occurrence of these animals on the surface of
the sediment may be correlated with oviposition since at
the time of collection (July) egg masses were present.
The sediment at this station is composed of sand with a
high silt/clay content. Eel Grass (Zostera marina Lin-
NAEUS, 1758) is the dominant organism. The specimens
examined from Duxbury, Massachusetts were apparently
collected from tidal flats but one lot came from “Eel
Grass Roots”.
SHELL
The shell is minute. The range in maximum length and
breadth is 1.0 - 1.86 mm and 0.74 - 1.40 mm respectively.
The majority of specimens is shiny and pellucid although
some have a chalky, opaque shell. The shell is ovate
with a broad mouth, with maximum breadth near the
middle of the shell. The average ratio of breadth to length
of all shells examined is 0.72. The lip of the aperture
does not extend above the apex. The rounded spire is
visible or invisible depending on how the shell is oriented
to the observer (Figures 1, 2). The sculpture of Philine
sinuata consists of a series of very fine concentric growth
lines of which some are stronger than others. A character-
istic of all specimens examined is a prominent growth
cessation line most clearly evident from the dorsal surface
(Figure 3).
Philine sinuata Stimpson, 1850
Figure 1
The right side of the sheil, as seen in this view, is tilted slightly
revealing the spire (Scale = 1mm)
Figure 2
In this view, the shell has been permitted to assume its normal
position on a flat surface (Scale = 1 mm)
Figure 3
Dorsal view of shell showing the spire and the prominent
growth cessation line
ANIMAL
The animal is translucent, yellowish in color with numerous
clusters of white pigment spots on the body, head shield
and epipodia. The buccal mass is red and is visible ex-
ternally through the epidermis. In the living animal, the
head shield is subtriangular and bluntly pointed posteri-
orly (Figure 4). This is not evident in preserved slugs.
The most characteristic external feature of the species is
the finger-like notch located in the midline of the pallial
margin posteriorly (Figures 4, 5a). A deep pallial notch
such as this also occurs in Philine pruinosa (CLARK, 1827).
RADULA
The radula consists of 9 to 16 rows, each with the formula
2-1-0°1-2. An admedian tooth (two views) and a pair
of lateral teeth are shown in Figures 6 to 8. The denticles
of the admedian teeth extend ventrally almost all the
way to the end of the blade. The blade then slants ab-
ruptly back to the frontal margin of the tooth. The
number of denticles varies from 14 to 25. Philine sinuata
does not possess jaw plates.
THE VELIGER
Vol. 12; No. 1
Philine sinuata Stimpson, 1850
Figure 4
Living animal, dorsal view (Scale = 1mm)
Figure 5
Living animal, right lateral view
a — pallial notch b-~ gill
(Scale = 1 mm)
c — cephalic shield d — foot
TAXONOMY
An unsuccessful search for the holotype of Philine sinuata
was made and we assume that it has been lost or de-
stroyed. However, the identity of Massachusetts and Con-
necticut animals with P sinuata offers no difficulties. The
general appearance of the animal is remarkably similar
to Stimpson’s description. The figure of the shell in
Stmmpson (1851) indicates a conspicuous, rounded spire
and a slight protrusion of the upper lip of the shell.
This apparent character led several European workers
to an incorrect conclusion as regards the identity of this
species with the European P denticulata (ApaAms, 1800).
Upon re-examination of the New England species, it is
now obvious that the appearance of the shell with regard
both to the spire and the protruding upper lip depends
Vol. 12; No. 1
THE VELIGER
Page 71
Radula of Philine sinuata Stimpson, 1850
Figure 6
A — left postero-lateral view of an admedian tooth (Scale = 10p)
Figure 7
A pair of lateral teeth (Scale = 10)
Figure 8
Right postero-lateral view of an admedian tooth (Scale = 10)
on the orientation of the shell to the observer. When
the shell is permitted to assume its natural position, aper-
ture up, on a flat surface, it appears as in Figure 2, i.e.,
the spire is not visible and the upper lip is acutely rounded.
If the outer lip is tilted up slightly, the rounded spire
becomes visible and the upper margin of the aperture
gives the appearance of being acute or keeled (Figure 1).
JEFFREYS (1867) and G. O. Sars (1878), both with some
hesitation, considered the European P. nitida (JEFFREYS,
1867) to be conspecific with P sinuata. ODHNER (1907)
concurred in this opinion. Subsequently, LEmcHeE (1948)
has shown that P nitida is identical with the older P
denticulata (ApAMS). It is now evident that these earlier
workers were incorrect in assigning P sinuata as a syno-
nym of P nitida (= P denticulata). Philine denticulata
is now well-known (Lemcue, 1948; Hortkxosu1, 1967)
and has, in addition to an acute and keeled apertural lip,
a radula formula 1:1:0-1-1. Furthermore, it lacks the
digital pallial margin of P sinuata.
Philine sinuata shares some of the characteristics of
the European P ventrosa (JEFFREYS, 1867). However,
in a recent publication LEmcHE (1967) has shown that
P. ventrosa (JEFFREYS, 1867) (= P velutinoides (G. O.
Sars, 1878) ) must be referred to an entirely new genus,
Rhinodiaphana. Consequently, P sinuata cannot be con-
fused with any European species. The American species
P. amabilis (VERRILL, 1880) is also a smooth shelled spe-
cies but has calcareous gastric plates as well asa 1:1:0°1:1
formula. Philine tincta (VERRILL, 1882) is another smooth
shelled American species but is inadequately described
and has never been rediscovered.
To the best of our knowledge, Philine sinuata is known
only from Maine, Massachusetts and Connecticut -— a
remarkably restricted distribution. However, if its associ-
ation with Eel Grass is characteristic, this species may
prove to be more widespread than indicated by its pres-
ently known distribution.
LITERATURE CITED
Goutp, Aucustus AppISON
1870. Invertebrata of Massachusetts.
Boston ; 524 pp.; 28 plts.
Horixosu1. M.
1967. Reproduction, larval features and life history of Phil-
ine denticulata (J. Apams) (Mollusca-Tectibranchia)
Ophelia 4 (1): 43 - 84
JEFFREYS, JoHN Gwyn
1867. British conchology.
LEMCHE, HENNING
1948. Northern and arctic tectibranch gastropods, II. A revi-
sion of the cephalaspid species. Kgl. Danske Vidensk.
Selsk. Skr. (Biol. Skr.) 5 (3): 29 - 136
1967. Rhinodiaphana G.N. ventricosa (JEFFREYS, 1865) re-
described (Gastropoda Tectibranchiata). Sarsia 29: 207 - 214
Opuner, Nits HjJALMAR
1907. Northern and arctic invertebrates in the collection of
the Swedish State Museum (Riksmuseum). III. Opisthobranchia
and Pteropoda. K. Svens. Vetensk. Akad. Handl. 41: 1 - 114
Sars, G. O.
1878. Bidrag til Kindskaben om Norges Arktiske Fauna I.
Mollusca regionis arcticae Norvegicae. Christiania
Stimpson, WILLIAM
1850. | Two new species of Philine obtained in Boston Harbor.
Proc. Boston Soc. Nat. Hist. 3: 333 - 334
1851. Shells of New England. A revision of the synonymy
of the testaceous mollusks of New England, with notes on
their structure, and their geographical and bathymetrical dis-
tribution. Boston
VERRILL, ADDISON E.
1880. Notice of recent additions to the marine invertebrata of
the North-East Coast of America, with descriptions of new
genera and species and critical remarks on others. Proc.
U.S. Nat. Mus. 3: 356 - 405
1882. Catalogue of marine Mollusca added to the fauna of
the New England region during the past ten years. Trans.
Conn. Acad. Arts & Sci. 5: 447 - 587
W. G. Binney (ed.),
Van Voorst, London, vol. 4
Page 72
THE VELIGER
Vol. 12; No. 1
Pleistocene Symbiosis:
Pinnotherid Crabs in Pelecypods from Cape Blanco, Oregon
BY
VICTOR A. ZULLO
AND
DUSTIN D. CHIVERS
California Academy of Sciences, San Francisco, California 94118
(Plate 5)
CoMMENSAL OR PARASITIC RELATIONSHIPS that are not
manifested by abnormalities in the preserved remains of
the host are difficult to establish in the fossil record. The
symbiont is often either without normally preservable
structures or is separated from its host during deposition
and burial. The discovery of the pinnotherid pea crab
Pinnixa faba (Dana, 1851) within paired valves of the
pelecypod Tresus capax (GouLp, 1850) from a late Pleis-
tocene terrace deposit at Cape Blanco, Oregon, is, there-
fore, noteworthy.
Extant adults of Pinnixa faba are found only in the
two Pacific Coast species of Tresus. In northern Califor-
nia where the range of the southern T: nuttalli (ConrapD,
1837) overlaps with that of the northern T: capax, P.
faba is found only in the latter (PEARcE, 1965, 1966).
Immature crabs, however, occur in a variety of pelecy-
pods, including species of Macoma, Mya, and Saxidomus.
The first fossil was discovered during preparation of a
general collection made in the Spring of 1960 for the
University of California Museum of Paleontology, Ber-
keley (UCMP locality A-8712). A second collection,
made specifically for additional pinnotherids in October,
1968, is deposited in the California Academy of Sciences,
Department of Geology (CAS locality 41270). Of 124
paired and presumably undisturbed shells of Tresus capax
examined, 15 contained reasonably complete crabs and
an additional 12 yielded fragments.
The pelecypods were found in living position near the
base of the Cape Blanco terrace deposit. Infaunal ele-
ments of this fauna, including Tresus capax, lived in a
shallow, offshore, sandy habitat below 5 fathoms (wave
base) under somewhat cooler hydroclimatic conditions
than those presently found off southern Oregon (Aopp1-
coTT, 1964). Fragments of molluscan shell dated both
by Carbon 14 and Thorium 230/Uranium 234 methods
indicate a minimum age of 33000 years for the deposit
(RicHarps & THURBER, 1966).
Although the size of pelecypods collected varied, all
but 3 of those yielding crabs are larger than 105 mm in
length. The 3 smallest shells are 82, 91 and 94mm, and
the largest 128mm, with 113mm being average size.
Sedimentary infilling of the shells varied greatly in coarse-
ness and degree of oxidation, and crabs were observed to
be most often preserved in finer-grained, unoxidized
fillings.
Extant individuals of Pinnixa faba are only slightly
calcified, and the fossils are accordingly delicate and dif-
ficult to extract from the unconsolidated sand filling. Each
pelecypod appears to contain but a single crab, as no
duplication of parts was observed. Elements of the cara-
pace, thorax, abdomen, and, rarely, articles of append-
ages are represented. The specimens are remarkably well
preserved. Thoracic musculature is evident in some frag-
ments (Plate 5, Figure 1), and in one individual the
optic region, including eyes, optic peduncle, epistome,
and suborbital region is present (Plate 5, Figure 2).
Even individual ommatidia can be discerned in this
specimen.
The majority of the specimens are females, although
some are too fragmentary for sexing. A single poorly pre-
served thorax is definitely that of a male, but it cannot
be identified to species (Plate 5, Figure 3). On the basis
of shape and sculpture of the carapace, and on the con-
figuration of the optic region, the identifiable remains
compare well with extant adult females of Pinnixa faba.
Pinnixa littoralis HotMEs, 1894, however, is also a com-
TuHE VELIGER, Vol. 12, No. 1 [ZuLLo & Curvers] Plate 5
Figure 1 Figure 2
Figure 1: Thoracic musculature in female Pinnixa sp. Hypotype
CAS 13207, width of thorax 9.2 mm
Figure 2: Optic region of female Pinnixa faba. Hypotype CAS
13208, width across eyes 4.8 mm
Figure 3: Thorax of male Pinnixa sp. Hypotype CAS 13209,
width 12mm
ad!
Vol. 12; No. 1
THE VELIGER
Page 73
mensal of Tresus, and it is possible that some unidentifiable
remains represent this species.
Traces of the leathery mantle and siphonal plates of
Tresus capax, together with ligament and partial mud
casts and molds of the gut and siphons are also preserved.
Although it was expected that the crabs would be found
near the anterior end of the shell where they might fall
after death, most occur near the middle, suggesting that
sedimentary infilling was rapid and contemporaneous
with tissue decay. Such rapid burial may well account
for the unusual preservation.
To our knowledge Pinnixa eocenica RaTHBUN, 1926,
from the late Eocene of southern Washington, and P
galliher. RatHBuN, 1932, PR montereyensis RATHBUN,
1932, and Parapinnixa miocenica RaTHBUN, 1932 from
the Miocene Monterey Formation at Pacific Grove, Cali-
fornia ate the only other pinnotherids to have been re-
ported from the Cenozoic of the Pacific Coast, but none
was found in association with a host (RaTHBUN, 1926,
1932). On the basis of the Pleistocene occurrence reported
herein, it appears likely that under similar condition of
preservation pinnotherids can be expected in Tertiary
and older pelecypod hosts.
RaTHBUN, Mary J.
LOCALITY DESCRIPTION
Localities UCMP A-8712 and CAS 41270 are in the SE
4 of the NE } of Sec. 2, T. 32S., R. 16W, Cape Blanco
quadrangle, Curry County, Oregon, near top of south
trending cliff south of road to Cape Blanco lighthouse
at 61 m elevation.
LITERATURE CITED
AppicoTT, WARREN O.
1964. _ A late Pleistocene invertebrate fauna from southwestern
Oregon. Journ. Paleont. 38 (4): 650-661; 3 figs.; pllts.
107 - 108
PEaRCcE, Jack B.
1965. On the distribution of Tresus nuttalli and Tresus capax
(Pelecypoda: Mactridae) in the waters of Puget Sound and the
San Juan Archipelago. The Veliger 7 (3): 166-170; plt.
27; 1 text fig. (1 January 1965)
1966. On Pinnixa faba and Pinnixa littoralis (Decapoda:
Pinnotheridae) symbiotic with the clam Tresus capax ( Pelecy-
poda: Mactridae). pp. 565-589 In: H. Barnes (ed.),
Some contemporary studies in marine science, George Allen &
Unwin, Ltd., London
1926. ‘The fossil stalk-eyed Crustacea of the Pacific slope of
North America.
pits. 1 - 39
1932. Fossil pinnotherids from the California Miocene.
U.S. Nat. Mus. Bull. 138: i - viii+ 1 - 155;
Journ.
Washington Acad. Sci. 22 (14): 411-413; 11 figs.
RicHarps, Horace G. « Davin L. THuRBER
1966. Pleistocene age determinations from California and Ore-
gon. Science 152: 1091 - 1092
Page 74
THE VELIGER
Vol. 12; No. 1
Escape Response of the Sea-Anemone
Anthopleura nigrescens (VERRILL) to its Predatory Eolid Nudibranch
Herviella BaBA spec. nov.
R. ROSIN
Department of Zoology, The Hebrew University of Jerusalem, Israel'
INTRODUCTION
EoLiD NUDIBRANCHS are generally associated with various
coelenterates on whom they feed, and whose non-everted
nematocysts they store. The eolids have evolved many dif-
ferent adaptations to their coelenterate prey (HyMaN,
1967). But coelenterates are not always a passive prey in
the association.
Swimming in the anemones Stomphia coccinea MULLER
and Actinostola spec. nov. (Actinostolidae) in contact
with Aeolidia papillosa (LinnaEus, 1761) (Aecolidiidae)
has been reported as a possible escape response (Rosson,
1966). A similar response is also elicited in these ane-
mones through electrical stimulation, contact with certain
sea stars, and in Actinostola spec. nov. through contact
with S. coccinea, but not vice versa (YENTSCH & PIERCE,
1955; Sunp, 1958; Ross & Sutton, 1964a, 1964b, 1967;
Warp, 1965; Rosson, 1966; Ross, 1967). Substances of a
different chemical nature, extractable from the sea star
and the eolid, cause swimming in the anemones, but no
chemical could be demonstrated in the response of one
species of anemone to the other.
The significance of the various swimming responses of
these anemones is not known. These actinostolids are deep-
sea species, at present not accessible to field work; there-
fore no information is available on their predator-prey
relationships in nature. The sea stars apparently do not
prey on these anemones. Rosson (1966) considers the
anemones’ response to the eolid an escape response be-
cause the geographic distribution of both is broadly re-
lated, and in the aquarium Aeolidia papillosa will bite
into these anemones. Ross (1967) states that the eolid
and the anemones are not frequently found living close
together in the field. Aeolidia papillosa, however, is not
« Present address: Beit-Oved, Israel
restricted as to depth. It is common in the intertidal area
and information is readily available on its feeding habits
there, where it is known to feed on a variety of anemones
and even hydroids (StEHOUWER, 1952; BRAAMS & GEELEN,
1953; Mixter, 1961). That is, it is not at all monophagous.
The fact that it is not frequently found near the actino-
stolids in the field may be due to escape of the anemones
from the vicinity of the eolid. At the same time, the fact
that the eolid will bite into the actinostolids in the aquari-
um may be an attempt of a polyphagous predator to
feed on prey with which it has no contact in nature.
The present is a report on the anemone Anthopleura
nigrescens (VERRILL, 1928) (Actinidae) and the eolid
Herviella Baza spec. nov. (Favorinidae). It is hoped that
it will shed further light also on the previous case.
ANIMALS anv HABITAT
Herviella spec. nov. mimics Anthopleura nigrescens and
can be mistaken for the anemone. Both species were found
in the same habitat in the intertidal area on a narrow strip
of beach near Kewalo Basin, Honolulu, Hawaii. The
anemones were abundant on loose rocks which lie on a
bottom of mixed sand and gravel, often exposed during
low tide. The column is equipped with suckers which at-
tach an “armor” of gravel and other foreign material.
When above water, the anemones close and contract, and
are thus completely shielded by the “armor”.
The eolids were found between February and August,
1966, mostly in pairs or larger groups, on the underside of
rocks with or without anemones. During a brief inspec-
tion in the field at night eolids were also observed crawling
above the rocks, and an eolid was encountered in the
midst of preying on a specimen of Anthopleura nigres-
cens which had its pedal disk detached from the rock.
Vol. 12; No. 1
Provisions for extensive field work at night could not be
made at the time. Eolids were not observed to feed in
the field during the day. Like the anemones, eolids were
often found above water level. They reproduced through-
out the period noted above. A few tiny specimens, as small
as 7mm long compared to up to 40 mm in adults, were
collected in June, indicating that occurrence in the inter-
tidal area is not restricted to the reproductive phase.
Neither species was found on the fringing reef away from
shore, or the sand between reef and shore.
LABORATORY OBSERVATIONS
In the aquarium eolids crawled up to water level when-
ever the air-pump was stopped ; they descended however to
feed on anemones attached to the floor of the aquarium,
They fed on the column and tentacles of Anthopleura
nigrescens, at night as well as during the day, but would
not feed on the only other unidentified anemone available
in the eolids’ natural habitat. The eolids approach A.
nigrescens, touch it repeatedly with the tentacles, bite and
attach with the jaws.
Anemones collected in the field and allowed to attach
to the aquarium floor were found to be detached from
the substratum when being preyed upon by the eolids.
But when it was noticed that (a) detachment of the
anemone occurred too rapidly to be attributable to de-
struction of the attachment apparatus; (b) a preying
eolid does not remain constantly attached to its prey, but
separates from the anemone every few minutes, and then
reattaches to it; (c) an anemone which detached from
the substratum due to eolid predation, reattaches readily
when removed from its predator; and when this sequence
of events was visualized under natural conditions, the
possible significance became apparent.
Therefore anemones were brought into the aquarium
still attached to the original rocks on which they had
been found in the field, and eolids were allowed to feed
on them. It was found that when an eolid attaches to an
anemone, the anemone responds by gradually shedding
the “armor” and detaching the pedal disc, all in a few
minutes. The eolid may lift the front part of its body and
carry the detached anemone upwards with it. For some
reason, eolid and anemone eventually separate. The re-
leased anemone rolls or drops off the rock to the floor
of the aquarium, where it remains lying on the column.
The fall is often interrupted by the falling anemone
attaching its tentacles to the rock, and sometimes even
settling on the pedal disk, or crawling on the column, then
detaching and resuming the downfall.
THE VELIGER
Page 75
Anemones which dropped off the rock, right them-
selves and attach to the floor of the aquarium in a few
minutes to half an hour at the most. The eolid on its part
very soon turns to feed on another anemone in its vicinity.
Thus, the intermittent feeding of an eolid on a single
anemone, which occurs with anemones attached to the
flat horizontal floor of the aquarium, inevitably trans-
forms into successive feeding on several different anem-
ones when these are attached to their natural, irregularly
shaped substratum.
Cases of partial detachment of the pedal disk, where
the anemone reattached on the same spot, without de-
taching all of the pedal disk, as well as cases of delayed
detachment only after separation from the predator, also
occurred. A single case was observed where an anemone
crawled on the pedal disk up the rock and away from the
eolid before it detached. Out of 67 recorded observations
on anemones bitten by an eolid, 66 detached within
minutes. In the one case which did not, the eolid fed for
over an hour on the tips of the tentacles only, and the
anemone shed part of its “armor”.
The exact mechanism of detachment could not be
determined by simple observations. However, in a few
cases, carefully observed, it was found that detachment
began by contraction of the pedal disk and column and
shedding of “armor” on the side attacked by the eolid,
and uplifting of the pedal disk on the opposite side,
with the anemone eventually tilting and falling away from
the eolid.
It is not known yet what is the direct stimulus for
shedding the “armor” and detachment of the pedal disk
of Anthopleura nigrescens preyed upon by Herviella spec.
nov. The anemones do not detach to simple mechan-
ical injury caused by prodding stainless-steel needles
through the column.
To lend further support to an interpretation of the
response of Anthopleura nigrescens to Herviella spec. nov.
as an escape response, it was important to assess the
survival value of the response. Therefore, a simple test
was conducted: three groups of 10 anemones each (a),
(b), (c) were obtained and placed in three small, sepa-
rate, equal aquaria respectively. Groups (a) and (b)
were obtained by mechanically scraping anemones off a
rock, taking care not to include specimens with injured
pedal disk due to scraping. Group (c) was obtained by
allowing eolids to feed on anemones attached to the rock,
and collecting anemones which escaped. Subsequently, 10
eolids were added to group (a) to assess maximum pos-
sible damage due to predation when ultimate escape is
not possible. Group (b) was kept as controls, which had
not been, and were not subsequently, subject to eolid
attack at all. Group (c) was kept to assess possible long-
Page 76
term effects induced by eolids prior to escape. The an-
swer to these simple questions is not apparent a priori,
because no one had studied them in these species. The
test was terminated at the end of 17 days.
It was found that in group (a) the eolids gradually
devoured and completely destroyed the anemones in 7
days; in groups (b) and (c) all anemones survived in
seemingly perfectly good condition.
This simple test shows that predation without escape
may proceed to the extent of total destruction of the
anemones, whereas anemones which escape are as per-
fectly viable as controls which had not been attacked
at all.
DISCUSSION
A natural predator-prey bond between the anemone An-
thopleura nigrescens and the eolid Herviella spec. nov. is
indirectly indicated by: 1. occurrence of both in a very
limited area in the same habitat; 2. a remarkable mimicry
between the two; 3. the occurrence of young specimens
of the eolid in the same habitat; 4. the fact that in the
aquarium Herviella spec. nov. feeds on A. nigrescens to
complete destruction of the anemone; it does not feed
on, or mimic, the only other anemone available in the
same restricted area; and eolids driven to the surface
by cutting off the air supply will descend to feed on A.
nigrescens attached to the aquarium floor. Also a specimen
of Herviella spec. nov. was actually encountered in the
field at night, with its mouth attached to a specimen of
A, nigrescens.
The detachment of the anemone to predation by the
eolid is considered an escape response in view of the
survival value of the response in the laboratory under
conditions simulating those in the field. The Anthopleura
nigrescens which was preyed upon by Herviella spec. nov.
in the field also had its pedal disk detached from the
rock.
The separation of predator and prey, which is an
essential link in the sequence of events which result in
escape of the anemone, is possibly due to slow writhing
on the part of the anemone, or to the eolid’s manner of
feeding.
Anthopleura nigrescens which detaches from its rock in
nature cannot attach to the loose bottom sand, and
water movement, not involved in the aquarium, is bound
to play a role in the intertidal area. In fact, a detached
A. nigrescens was encountered in the field, lying on the
sand above water level at night, as well as one being
carried by surf during the day. This is not to suggest
that their detachment was necessarily due to eolid pre-
dation. Nor is it being suggested that escape from its
THE VELIGER
Vol. 12; No. 1
predatory eolid, Herviella spec. nov., is the sole function
of the ability of A. nigrescens to shed foreign material
attached to its suckers, and detach the pedal disk.
The variety of adaptations of eolids to their coelen-
terate prey suggests a very ancient association between
these two groups. The response of Anthopleura nigrescens
to Herviella exhibits certain similarities to the response of
the Actinostolidae to A. papillosa. It is perhaps too early
to say whether these similarities are superficial only, or
not. The eolids belong to two different families, whereas
the Actiniidae belong to the subtribus Endomyaria, and the
Actinostolidae to the Mesomyaria. This raises the question
whether these are two isolated cases. It may be worthwhile
noting that Tarpy (1962, 1965), who was interested in
eolids but paid little attention to the behavior of their
coelenterate prey, notes that anemones preyed upon by
several eolids he was studying, finally detached.
ACKNOWLEDGMENTS
The case was found during studies on a scholarship of
the National Council for Research and Development,
Government of Israel. I thank Dr. C. E. Cutress of the
University of Puerto Rico for identification of the ane-
mone, and Dr. E. A. Kay of the University of Hawaii
for pending identification of the eolid.
LITERATURE CITED
Braams, W.G., & H. F. M. GEELEN
1953. The preference of some nudibranch eolids for certain
coelenterates. Arch. Néerl. Zool. 10: 241 - 264
Hyman, Lippy HENRIETTA
1967. The invertebrates.
249 text figs.
MILLER, MIcHAEL C.
1961. Distribution and food of the nudibranchiate Mollusca
of the South of the Isle of Man. Journ. Anim. Ecol. 30:
95 - 116
Rosson, ELAINE A.
1966. Swimming in Actiniaria.
McGraw Hill Inc. 6: vii+ 792 pp.;
Symp. Zool. Soc. London
16: 333 - 360
Ross, D. M.
1967. Behavioural and ecological relationships between sea
anemones and other invertebrates.
Ann. Rev. 5: 291 - 316
Ross, D. M. « L. Sutton
1964a. The swimming response of the sea-anemone Stomphia
coccinea to electrical stimulation. Jounr. Exp. Biol. 41:
735 - 750
1964b. Inhibition of the swimming response by food and of
nematocyst discharge during swimming in the sea anemone
Stomphia coccinea. Journ. Exp. Biol. 41: 751 - 758
1967. _ Swimming sea anemones of Puget Sound: swimming of
Actinostola n. sp. in response to Stomphia coccinea. Science
155; 1419 - 1421
Oceanogr. Mar. Biol.
Vol. 12; No. 1
THE VELIGER
SrEHOUWER, E. C.
1952. The preference of the slug Aeolidia papillosa (L.)
for the sea anemone Metridium senile (L.). Arch.
Néerl. Zool. 10: 161 - 170.
Sunp, P.
1958. A study of the muscular anatomy and swimming behav-
iour of the sea anemone Stomphia coccinea. Quart. Journ.
micr. Sci. 99: 401 - 420
Tarpy, J.
1962. A propos des espéces de Berghia (Gastéropodes, Nudi-
branches) des cétes de France et leur biologie. Bull. Inst.
Oceanogr. Monaco 1255: 1 - 20
1965. Description et biologie de Cerberiella bernadetti, espéce
nouvelle de gasteropode nudibranche de la c6te atiantique fran-
Gaise. Bull. Inst. Oceanogr. Monaco 1349: 1 - 22
Waprp, J.
1965. ‘An investigation on the swimming reaction of the ane-
mone Stomphia coccinea. I. Partial isolation of a reacting
substance from the asteroid Dermasterias imbricata. Journ.
Exp. Zool. 158: 357 - 364
YENTCH, CHARLES SAMUEL & D. C. PIERCE
1955. A “swimming” anemone from Puget Sound. Science
122: 1231 - 1233
Page 77
Page 78
THE VELIGER
Vol. 12; No. 1
A Three-Dimensional Representation of Measurement Data
BY
PAUL CHANLEY
AND
W. A. VAN ENGEL
Virginia Institute of Marine Science, Gloucester Point, Virginia 23062 '
(4 Text figures)
IT HAS BEEN FOUND convenient in describing bivalve lar-
vae to present length, height and depth relationships in
a three-dimensional graph (CHANLEy, 1969). Such a
graph may also be used to illustrate other kinds of meas-
urement data. Dimensions of the larvae of a small com-
mensal bivalve, Montacuta percompressa Dauu, 1899,
have been plotted to illustrate the construction and inter-
pretation of the graph.
The vertical axis represents height and the horizontal
axis depth (Figure 1). The axis at 45° between these two
represents length. The vertical plane formed from the
height axis along the length axis and the horizontal plane
formed from the depth axis along the length axis provide
surfaces on which two-dimensional representations of the
height : length and length : depth relationships can be
plotted. Height and depth coordinates extend from their
respective axes parallel to both the height and depth axes
and form right angles at the length axis. The line AB
represents a height of 1694; EH a depth of 77; and
IJL a length of 200u. Length 200u by height 169 is
marked where lines AB and IJ intersect, length 200u by
depth 77 where lines EH and IL intersect. Measure-
ments of larval Montacuta percompressa have been plot-
ted on the appropriate planes as dots which have been
enclosed with lines by inspection. Line MN represents
minimum depth, OP maximum depth, QR minimum
height, and ST maximum height for any length.
A three-dimensional concept of the length :height:depth
relationship can now be advanced by considering each
measurement a plane. For example, height 169 that was
represented as line AB in Figure | may also be represented
as plane ABCD in Figure 2. Similarly lines EH and IJL
can be represented as planes EFGH and IJKL, respec-
« Contribution No. 311, Virginia Institute of Marine Science
tively. The broken lines indicate the intersections of these
planes; length 200u by height 1694 by depth 77p is
the point W, located at the intersection of all three planes.
Point W is one vertex of a base plane that defines the
limits of height and depth for a length of 200u. The
other vertices of the base plane can be obtained (1) by
finding the points where the length plane for 200u (IJKL)
intersects lines MN, OP, OR, and ST (Figure 3), and
(2) extending horizontal and vertical lines (broken) from
these points to new points of intersection W, X, Y and Z.
Thus, at length 2004, point W represents the minimum
height and depth, X maximum height and minimum
depth, Y maximum height and depth, and Z minimum
height and maximum depth. Lines WX and YZ equal
the range in height for length 200, while lines WZ and
XY equal the range in depth. Base plane WXYZ encom-
passes all possible combinations of height and depth for
a length of 200u.
Similar base planes may be constructed at selected
lengths (as shown for 250p in Figure 3) over the entire
length range. When the vertices of all base planes are
connected by lines, the resulting three-dimensional figure
(Figure 4) encompasses all possible length :height :depth
combinations. This figure is a polyhedron consisting of a
series of rectangular oblique prisms whose bases are
parallel planes.
If the polyhedron overlaps the length axis or is other-
wise poorly placed, it can be raised or lowered by adding
or subtracting units at the base of the height axis. It can
be moved laterally or backward and forward by similar
manipulation of the depth and length axes.
Construction of a polyhedron is a relatively easy
method of reducing a series of 3 continuous variables to
a simpler form. Because the data are not reduced to
statistics to which tests of significance can be applied,
Vol. 12; No. 1
THE VELIGER
Page 79
Two-dimensional graphs of height:length and length:depth
relationships shown in a three-dimensional diagram, for larval
Figure 1
Montacuta percompressa DAuL, 1899. Dots represent actual obser-
vations. Lines MN, OP, OR, and ST represent minimum and
maximum limits on length and depth.
Height
as
o°8
33
ay
Py
4S
3 y
*
ey
275
250
225
200
© ° 9
NG ) 2 ¥
Depth
300
325
375
Page 80 THE VELIGER Vol. 12; No. 1
Figure 2
Three-dimensional diagram of plane surfaces representing height at
169 », depth at 77 », and length at 200 ». Broken lines represent the
intersection of pairs of planes. W is the point of intersection of
all three planes.
Vol. 12; No. 1 THE VELIGER Page 81
Height
Figure 3
Three-dimensional diagram showing the construction of base planes
WXYZ and 1, 2, 3, 4, utilizing data from the two-dimensional
graphs of height:length and length:depth. WXYZ and 5 OH Gy fh
encompass all height and depth combinations for lengths of 200 p
and 250 p, respectively.
fa,
So
‘8,
2%
Page 82
THE VELIGER
Vol. 12; No. 1
Height
“ee
So
%
90
“25
So
(o)
Figure 4
ey f :
> Three-dimensional diagram showing completed polyhedron
encompassing all length: height:depth combinations of larval
Montacuta percompressa and two-dimensional graphs of height:
length and length:depth. Base planes are shown for length 115»
and at 25 intervals of length from 150 to 375 p.
Vol. 12; No. 1
THE VELIGER
comparisons with other sets of measurements are limited.
An alternative method, multiple regression analysis, would
provide a meaningful statistical statement, but could not
be used if the variates do not follow a multivariate normal
distribution where linearity of regression is requisite.
Each set of measurement data for any species of
bivalve larvae may havea characteristic polyhedron graph-
ically distinguishable from other polyhedra. Similarly,
averages and ranges may be peculiar to each set. If
polyhedrons are determined for a sufficient number of
species computer identification of planktonic larvae may
be possible.
We would like to express our appreciation to Mrs. Jane
S. Davis for preparing the illustrations necessary for this
paper.
LITERATURE CITED
CHANLEY, PAUL
1969. Larval development of the coquina clam, Donax vari-
abilis, Say, with a discussion of larval hinge structure in the
Tellinacea. Bull. Mar. Sci. (in press)
Page 83
Page 84
THE VELIGER
Vol. 12; No. 1
Notes on the Mitridae of the Eastern Pacific II]
The Genus Thala, with the Description of a New Species
BY
GALE G. SPHON
Los Angeles County Museum of Natural History, Los Angeles, California 90007
(Plate 6; 2 Text figures; 1 Map)
AMONG THE ALLAN Hancock FouNnpATIoN gastropods
that are on loan to the Los Angeles County Museum of
Natural History and in material at the California Aca-
demy of Sciences is an undescribed species of the family
Mitridae from the Galapagos Islands. The morphological
characteristics of this form indicate that it should be
placed in the genus Thala H. & A. ApaMs, 1853. I would
also place two other Eastern Pacific species, Mitra grati-
osa REEVE, 1845, and M. solitaria C. B. Apams, 1852, in
Thala.
Thala was used for some Eastern Pacific members of
the Mitridae (Dati, 1921; Oxtproyp, 1927), but was
dropped in favor of Mitromica Berry, 1958. However,
I regard the latter as a synonym for reasons that are
discussed later in this paper.
Thala H. « A. Apams, 1853
Thala H.« A. Apams, 1853, p. 178 (as a subgenus of Mitra).
Type species by S.D., CossMann, 1889: Mitra mirifica
Reeve, 1845a. Recent, Capul, Philippine Islands. REEVE,
1845b, Conch. Icon. plt. 34, spec. 277
Micromitra BELvarpi, 1888, p. 147. Type species by S.D.,
Coan, 1966, Micromitra taurina BeLuarpi, 1888. Middle
Miocene, Europe
Mitromica Berry, 1958, p. 94. Type species by O. D., Mitra
solitaria C. B. ApaMs, 1852
DISCUSSION
Thala is a genus of small species that can very roughly
be divided into 3 groups, but the groups are far too indis-
tinct to form subgenera. The first group has a recurved
canal and includes the type species T: mirifica (REEVE,
1845) and such species as T: recurva (Reeve, 1845) and
T. todilla (Micuets, 1845). Unfortunately, the radula of
no member of this group has been illustrated.
The second group is intermediate, with a slightly re-
curved canal; it includes such species as Thala cernica
(Sowerby, 1874), T. milium (Rutve, 1845), and T. soli-
taria (C. B. Apams, 1852).
The third group has a more truncate canal and includes
Thala floridana (Daut, 1884), T. gratiosa (REEVE, 1845),
and T: ogasawarana (Pitssry, 1904). All the radulae of
the genus Thala that have been figured (McLean, 1967;
Crernonorsky, 1966; Hane, 1943; and Tues, 1931)
seem to belong to this group.
Berry’s diagnosis of his taxon, Mitromica, was based
on what is actually Thala gratiosa although he cited as the
type species Mitra solitaria. Mitromica must remain with
the species named as type, whatever BErry’s concept
of Mitra solitaria.
THIELE (1931) considered Thala to be a subgenus of
Pusia, based on the 3 prominent cusps of the rachidian
plate and the sickle-shaped lateral plates. However, Thala
has several morphological features not shared by Pusza.
The most significant of these are the prominence of the
lirations on the inner side of the labrum, the presence or
suggestion of an anal sulcus, cancellate sculpture, and a
subsutural band. These characters, as well as some features
of the radula, make a distinguishable genus.
Both Daty (1921) and Otproyp (1927) listed Mitra
orcutti Dat, 1920, in the “section” Thala of the genus
Mitra. However, McLEan (personal communication) has
examined the holotype and finds the species to be a syno-
nym of the turrid Mitromorpha gracilior (Tryon, 1884).
Thala gratiosa (Reeve, 1845)
(Plate 6, Figures 1, 2)
Mitra gratiosa Reeve. 1845, p. 53
Thala gratiosa (REEVE, 1845). - Tryon, 1882, p. 161
Mitra (Thala) solitaria C.B. Apams, 1852. - Datt, 1921, p.
87. - Oxproyp, 1927, p. 173. - Smiru, 1944, p. 33
Vol. 12; No. 1
Mitromica solitaria (C. B. ApaMs, 1852). - Berry, 1958, p.
44. - McLean, 1967, p. 58, text fig. 1 (radula)
Mitra (Mitromica) solitaria C. B. ADAMS, 1852. - Keen, 1958,
p. 428
Mitra (? Costellaria) nodocancellata Stearns, 1890, p. 213
Diagnosis: Shell small for the genus, adult specimens to
10 mm in length; black; ovate; sculpture cancellate; canal
truncate; aperture narrow; labrum lirate within; anal
sulcus becomes more pronounced with age.
Type Material: The holotype of Mitra gratiosa REEVE,
1845, is in the British Museum (Natural History). Type
Locality: Six fathoms, Galapagos Islands, Ecuador, Hugh
Cuming, collector. The holotype of Mitra nodocancellata
Stearns, 1890, is in the United States National Museum
(no. 55490). Type Locality: Gulf of California, W. J.
Fisher, collector.
Distribution: Thala gratiosa has been reported as Mitra
solitaria from Point Loma, San Diego, California (Dat,
1921; Oxproyp, 1927; SmirH, 1944; Keen, 1937). Ap-
parently this is the northern limit of its distribution and it
is rare in the San Diego area, for it has not been reported
from there in over 30 years. It is unknown between San
Diego and the Cape San Lucas area in Baja California.
The species is found throughout the Gulf of California
and south to Panama and the Galapagos Islands.
I have examined approximately 300 specimens of this
species and while I do not consider it rare, it certainly
is not a common species.
Discussion: Dati (1921) was apparently unaware of
Thala gratiosa from the Galapagos Islands when he placed
Mitra nodocancellata in synonymy with T. solitaria, be-
cause he cites the photograph of STEARNS’ (1890) holo-
type as an example of T. solitaria. Apparently this error
has been perpetuated for 2 reasons: 1) ‘Thala gratiosa
was originally described from the Galapagos Islands and
never reported from the mainland, and 2) it was de-
scribed as having a brown shell while T. solitaria was
described as having a black one. This would indicate
that Cuming had dredged a dead specimen, for T: gratiosa
usually bleaches to a dark rust or brown color after the
animal has died.
Thala gratiosa has an Atlantic analogue in T. floridana
(Da, 1884). The main differences between the 2 spe-
cies, other than distribution, is that T: floridana is smaller
and proportionally wider.
Thala solitaria (C. B. ApAMs, 1852)
(Plate 6, Figure 3)
Mitra solitaria C.B. ADAMS, 1852, p. 44. - TuRNER, 1956, p.
87, plt. 5, fig. 1 (holotype)
THE VELIGER
Page 85
Thala solitana (C.B. Apams, 1852). - Tryon, 1882, p. 160,
pit. 47, fig. 358
Mitromica solitaria (C. B. ApamMs, 1852. - Berry, 1958, p. 44
(type designation only)
Mitra (Mitromica) solitaria C. B. ApAMs, 1852. - Keen, 1958,
p. 429, fig. 646 (figure on the right, after Turner, 1956)
Not “Mitra solitaria” of other authors
Diagnosis: Shell relatively large for the genus, adult spe-
cimens to 17 mm in length; black, occasionally with white
nodes; sub-attenuate; sculpture cancellate; canal slightly
recurved ; aperture moderately narrow; labrum lirate with-
in; anal sulcus becoming more pronounced with size and
age; a subsutural band present.
Type Material: The holotype is in the Museum of Com-
parative Zoology at Harvard University, no. 186351. Type
Locality: Panama, C. B. Adams, collector.
Distribution: The present report on the distribution of
Thala solitaria is based on 10 specimens that show it to
occur from Los Arcos, Banderas Bay, Jalisco, Mexico, to
the Galapagos Islands.
With the exception of the holotype, one specimen from
Panama in the Santa Barbara Museum of Natural His-
tory (cat. no. 06291), and 2 specimens from the Galapa-
gos Islands in the California Academy of Sciences (loc.
no. 27221), all other specimens examined are in the Los
Angeles County Museum of Natural History.
Discussion: Thala solitaria is evidently rare and closely
related to T: gratiosa. As in that species empty shells
bleach to a dark rust or brown color. The 2 species have a
very similar habitat. ADAMS (1852) said that T: solitaria
was found “... Under stones near low water mark...”,
and when T: gratiosa is found intertidally it is always
found under rocks.
Thala jeancateae SPHON, spec. nov.
(Plate 6, Figure 4)
Diagnosis: Shell medium sized for the genus, white with
brown markings, sub-acuminate; sculpture cancellate;
canal slightly recurved; aperture moderately narrow; lab-
rum lirate within; anal sulcus and subsutural band present.
Description of Holotype: Shell medium sized for the
genus, length 9.4mm, width 3.8mm, length of aper-
ture 2.4 mm; sub-acuminate; whorls 9; nucleus and first
3 postnuclear whorls smooth; sculpture of the remaining
6 of sharply incised spiral and axial lines giving a nodose-
cancellate appearance; anal sulcus evident but faint;
labrum thin with denticulations within; columella with 4
strong plaits; anterior canal short, slightly recurved; sub-
sutural band set off by a row of nodose-cancellate bead-
Page 86
ing; color white with lines and irregular smudges of
brown, the base of a more intense brown; aperture, co-
lumella, and plaits porcelain-white.
Type Material: The holotype is in the Los Angeles County
Museum of Natural History Invertebrate Zoology Type
Collection (LACM-AHF 1202) and was dredged by the
Allan Hancock Pacific Expedition of 1934 on January
Figure 1
Radula of Thala jeancateae SPHoN, spec. nov.
The rachidian plate and one lateral plate
15, 1934, station no. 155-34. There is one paratype (CAS
13204) and one hypotype in the California Academy of
Sciences. The paratype is a dead specimen from the type
locality, but no depth is given. The hypotype is a Pleisto-
THE VELIGER
Vol. 12; No. 1
cene fossil from James Island in the Galapagos Islands
(CAS loc. no. 27255).
Type Locality: 50-60 fathoms off Tagus Cove, Aibe-
marle Island, Galapagos Islands, Ecuador (0°16'45”S,
SI 22/520)
Name: The name of this species, jeancateae, has been
chosen in honor of Jean-Cate (Mrs. Crawford N.) for
her work with the Mitridae.
Discussion: ‘Thala jeancateae can readily be distinguished
from the 2 other Eastern Pacific members of the genus, T:
gratiosa and T. solitaria, by the color. Thala jeancateae is
white with brown streaks and smudges, while T: gratiosa
and T: solitaria are black. 'Thala jeancateae differs in
shape from T.solitaria, being more sharply attenuate, and
from T: gratiosa in being acute rather than obtuse. The
blunt spire of T: gratiosa also serves to separate it from the
other 2 species which have sharp spires.
The sculpture on all 3 species is cancellate. The sculp-
ture in Thala jeancateae and T. solitaria is very similar but
differs from T. gratiosa in that both the spiral and axial
ribs are wider and more nodose at their juncture. In T:
gratiosa the sculpture is less nodose and the spaces be-
tween the ribs give the impression of forming squarish
pits.
The radulae clearly separate 2 of the 3 Eastern Pacific
species of ‘Thala. (Unfortunately, I have not been able to
obtain the radula of T: solitaria). 'Thala jeancateae has
the typical three-cusped rachidian plate (see Text figure
1) indicated by CERNoHoRSKY (1966, p. 120) for T.
ogasawarana (PitsBry, 1904) and T: simulans (von Mar-
Explanation of Plate 6
Figure 1
Mitra gratiosa REEVE, 1845
Holotype. Galapagos Islands, Ecuador. Length 11.5 mm
Photograph courtesy of British Museum (Natural History)
Figure 2
Mitra nodocancellata STrEaRNs, 1890
Holotype. Gulf of California, Mexico. Length 10mm
Photograph courtesy of Dr. James McLean, Los Angeles County
Museum of Natural History
Figure 3
Mitra solitaria C. B. Apams, 1852
Holotype. Panama. Length 0.68 inches
Photograph courtesy of Dr. Ruth Turner, Museum of Comparative
Zoology, Harvard University
Figure 4
Thala jeancateae SPHON, spec. nov.
Holotype. 50 - 60 fathoms off Tagus Cove, Albemarle Island,
Galapagos Islands, Ecuador. Length 9.4mm. Photograph courtesy
of Mr. Lawrence Reynolds, Los Angeles County Museum
of Natural History
Tue VELIGER, Vol. 12, No. 1 [SproNn] Plate 6
Figure 1 Figure 2 Figure 3
Vol. 12; No. 1 THE VELIGER Page 87
San Diego
Rancho £1 Tule
Carmen Island
Los Angeles Bay
San Luis Gonzaga Bay
San Luis Island
Puertecitos
Agua Chale
Guaymas
Mazatlan
Banderas Bay
Cuastocomate Bay
San Juan Del Sur KILOMETERS
* 0 50100 1000
Corinto ri
0 50 100 500
Bahia Honda STATUTE MILES
Balboa
Borgona Island
Port Utria
Barrington Island
Localities: Thala gratiosa (REEVE, 1845), stations 1 to 19
Thala jeancateae Spuon, spec. nov., station 19
Thala solitaria (C. B. ApaMs, 1852), stations 11, 12, 16-19
Page 88
Figure 2
Thala gratiosa (REEVE, 1845) (after McLEAN, 1967)
The rachidian plate and one lateral plate
TENS). Thala gratiosa (figured by McLean, 1967, as Mit-
romica solitaria) is more aberrant (see Text figure 2)
with 7 cusps on the rachidian plate.
ACKNOWLEDGMENTS
I am indebted to the many people and institutions who
allowed me to examine their collections of this group. I
am particularly indebted to Dr. Rudolf Stohler and Mrs.
Jean Cate for their help and understanding, Dr. James
McLean and Dr. Eugene Coan for critical reading of the
manuscript, and Mr. James Lance for his mounting and
illustration of the radula of Thala jeancateae.
LITERATURE CITED
ApaMs, CHARLES BAKER
1852. Catalogue of shells collected at Panama, with notes on
synonymy, station, and geographical distribution. Ann. Lyc.
Nat. Hist. New York 5: 229 - 344 (June) ; 345 - 548 (July)
ApaMs, Henry « ARTHUR ADAMS
1853. The genera of Recent Mollusca arranged according to
their organization. London (John van Voorst) 3: i-xl+
1 - 484
Bexxarot, Luict
1888-1889. I molluschi dei terreni Terziarii del Piemonte e
della Ligura. Part 5: Mitridae Mem. Acc. Sci. Torino (2)
38: 79-166; plts. 1-2; 257-326; plts. 3-4 [1888, possibly
1887]; (2) 39: 145-194; plts. 5-6 [1889, possibly 1888]. Also
issued separately.
THE VELIGER
Vol. 12; No. 1
Berry, SAMUEL STILLMAN
1958. | West American molluscan miscellany. - II.
in Malacol. 1 (16): 91-98
CERNOHORSKY, WALTER OLIVER
1966. A study of mitrid radulae and a tentative generic ar-
rangement of the family Mitridae. The Veliger 9 (2): 101
to 126; 47 text figs. (1 October 1966)
Coan, EucENE VICTOR
1966. | Nomenclatural units in the gastropod family Mitridae.
The Veliger 9 (2): 127 - 137 (1 October 1966)
CossmMaNnn, ALEXANDRE EpouARD MAUurRICE
1899. _ Essais de paléoconchologie comparée 3: 1 - 201; 8 plts.
Paris
Dati, WiLt1AM HEALEY
1921. Summary of the marine shellbearing mollusks of the
northwest coast of America, from San Diego, California, to
the Polar Sea... . Bull. U. S. Nat. Mus. 112: 1 - 217; plts.
1-22 (24 February 1921)
Hasse, TaDASHIGE
1943. On the radulae of Japanese marine gastropods (I).
Japan. Journ. Malac. 13 (1-4): 68-76; plts. 3-4
Keen, A. Myra
1937. An abridged check list and bibliography of West North
American marine mollusca. Stanford Univ. Press, Stanford,
Calif. pp. 1-88
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi + 624 pp.; illus.
Stanford Univ. Press, Stanford, Calif. (5 December 1958)
McLean, JAMES HAMILTON
1967. Note on the radula of Mitromica Brrry, 1958. The
Veliger 10 (1): 58; 1 text fig. (1 July 1967)
O.proyp, IpA SHEPARD
1927. The marine shells of the west coast of North America.
2 (1): 297 pp.; 22 plts. Stanford Univ. Press, Stanford, Calif.
ReEEvE, LovELL Aucustus
1845a. Descriptions of eighty-nine new species of Mitra, chiefly
from the collection of H. Cuming, Esq. Proc. Zool. Soc.
London 13: 45 - 61 (September 1845)
1845b. Conchologia Iconica. Monograph of the genus Mitra.
10: plts. 1 - 39
SmituH, MAxwELL
Leafl.
1944. | Panamic marine shells.
912 figs.
STEARNS, ROBERT EDWARDS CARTER
1890. Scientific results of explorations by the U.S. Fish Com-
mission steamer Albatross. No. XVII. - Descriptions of new
West American. land, fresh-water, and marine shells, with notes
and comments. Proc. U.S. Nat. Mus. 13: 205 - 225; plts.
15-17
THIELE, JOHANNES
1929. | Handbuch der systematischen Weichtierkunde.
Jena, Gustav Fischer, 1929-1935; 1- 1154; 893 text figs.
Tryon, GeorcE WASHINGTON, Jr.
1882. | Manual of Conchology. Monograph of the Mitridae.
4: 106 - 200; pits. 32 - 58. Philadelphia.
Winter Park, Florida, 127 pp.
Vol. 12; No. 1
THE VELIGER
Page 89
Cuttlebones on the Beach at Galveston
BY
HAROLD W. HARRY
AND
SELMA F SNIDER
Marine Laboratory, Texas A&M University, Galveston, Texas 77550
(3 Text figures)
“To him who in the love of nature holds
Communion with her visible forms, she speaks
A various language for his gayer hours
She has a voice of gladness, and a smile
And eloquence of beauty; and she glides
Into his darker musings with a mild
And healing sympathy, that steals away
Their sharpness ere he is awake.”
THESE LINES FROM William Cullen Bryant’s ‘““Thanatop-
sis’ are particularly appropriate to the pass-time of
beach combing, for no matter how barren a beach may
be, or how frequently one has combed it, there is always
a refreshing release from the cares of the day. Not in-
frequently, there is something new to excite the imagi-
nation, which may speak to each a “various language.”
One such occasion was experienced by the junior author
in January 1968. When combing the beach at 61" Street
in Galveston, she came upon a cuttlebone. Shortly she en-
countered Mrs. Rosalie Schreiber, who had picked up
six more and neglected several others. Neither person had
ever seen them on the beach before, although both have
been ardent beach combers here for many years. Inquiry
among devoted beach combers of the Galveston Shell
Club revealed only one other cuttlebone. Mrs. Louise
Beeman had found one some time previously. Her speci-
men is larger than the ones found by Mrs. Snider and Mrs.
Schreiber, and too worn to be diagnosed much more
specifically than to say that it is a cuttlebone.
Where did they come from? Are there cuttlefish (Sepia)
in the western Atlantic? If they floated here from some
distant shore, from which one, and are cuttlebones suffi-
ciently diagnostic to determine the species of cuttlefish?
Perhaps they were thrown here by someone who had
gotten them elsewhere? Very occasionally marine objects
not indigenous to this region are found on the beach at
Galveston, probably thrown there by some irresponsible
person. West coast abalone shells, also Cypraea moneta
and Melongena corona have on occasion been found, and
doubtlessly came from the shell shops along the sea wall.
The local pet shops sell cuttlebones, which pet owners
give to caged birds to sharpen their beaks. All that we
have seen in the pet shops are almost twice as long as the
ones found on the beach in January 1968. They were
moreover carefully trimmed with a knife along their lat-
eral margins and ventral surface. That the beach speci-
mens were casually discarded seems unlikely.
The margins of the beach specimens were not trimmed,
but the ventral surface was slightly worn. They had no
marine growths (barnacles, etc.) on them, and all had a
strong odor recalling that peculiar to spoiled squid. This
suggests that they did not float in the sea very long.
But Sepia is reputedly absent from the western Atlantic,
and indeed all shores of the American continents (HoyLe,
1886, p. 223; Lane, 1962, p. 29). Several experienced
shrimp fishermen of Galveston to whom we showed a pic-
ture of the cuttlefish said that they had never seen it in
these waters. We have not found cuttlebones listed in
any of the numerous faunal lists of mollusks based chiefly
on shore collecting in the western Atlantic. Such lists are
not very critical in differentiating between species which
actually live in an area, and those which are adventitious,
for they frequently list Spzrula, a shell common on beaches,
but known to live only between 200 and 1500 m depth
(BruuN, 1943). In 1968, the senior author found no
Page 90
cuttlebones in the collection of the U.S. National Muse-
um from beaches along the shores of the Americas, but
beach specimens from other shores were well represented.
Along the beaches of England, North Africa, and the
Indian Ocean, cuttlebones are said to be common. ForBES
& Haney (1853, vol. 4, p. 238) noted that “The common
cuttlefish (Sepia officinalis L.) is one of the most beauti-
ful and curious of British mollusks, but although its bone,
or shell, is frequently cast up on all our sandy shores, the
creature itself is rarely seen and seldom taken.” Lane
(1962) noted that tons of cuttlebones are exported yearly
from Tunisia, apparently gathered from the beach, al-
though he is not explicit. Of the numerous new species of
Sepia named since Hoye (1886) summarized the known
Recent Cephalopoda, several have been described from
cuttlebones cast upon beaches. Thus, SmirH (1916) de-
scribed four new species of Sepia from beach cuttlebones
in South Africa, and WinckworTH (1936) one species,
based on two cuttlebones found at Madras, India.
There is one recent report of beach cuttlebones in the
western Atlantic. ErpMAN (1957) reported finding several
at Anegada, the northeasternmost of the British Virgin
Islands. They were found high up on the beach, had been
worn by the sea, and “one or two had a greenish tinge in
cross section, indicating perhaps some algal growth.” One
specimen was sent to Dr. Gilbert Voss, who identified it
as Sepia officinalis. Dr. Voss suggested that it had floated
across the Atlantic, and noted that he had several speci-
mens from the east coast of Florida, all of which showed
evidence of being at sea for a long time. ERDMAN noted
that he had never found cuttlebones in Puerto Rico,
where he has spent much time beach combing.
A very early report of Sepia in the Caribbean has been
discounted by Hoyte (1886) as too vague to be useful.
But, as p’OrBIGNY was an astute observer of mollusks,
and a specialist in cephalopods, his statement on the
matter is worth quoting in full (p’Orsicny, 1841, p. 33,
translated from the French) :
“No. 8. Cuttlefish of the Antilles
“Sepia antillarum d’Orb.
We designate, under this purely provisional name,
the cuttlefish indicated by Browne [footnote: The
natural history of Jamaica, p. 386] as occurring at
Jamaica. It is rather common at the other Antilles,
and we have before our eyes an example from Mar-
tinique preserved in the Museum of Paris, under the
improper name of Sepia Orbigniana Fer. After
having examined it with care, although someone had
_ removed the ossicle from it, it is easy for us to recog-
nize that this is not Sepia Orbigniana, but indeed a
species one no longer may place near Sepia vulgaris,
from which by the poor state of preservation and the
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Vol. 12; No. 1
absence of the ossicle we are prevented from recog-
nizing the identity or the specific differences.
“However, we do not believe that we should pass in
silence this (p. 34) species of the Antilles, but in
order to arrive at a better understanding of it, arouse
attention in this respect.”
We have not seen thie rare book by BRowNE (1756)
which p’OrBicNy cites as recording the presence of the
cuttlefish in Jamaica. Nor can we account for p’Orsic-
Ny’s comparison of Antillean cuttlefish with Sepia vul-
garis, for no such trivial name seems to have been used in
the genus since 1758. Browne seems not to have used
the name. Mr. Thomas H. Farr of the Science Museum
of the Institute of Jamaica, sent a copy of BROowNeE’s de-
scription, evidently taken from the second edition. BROWNE
did not give an illustration of it. The description (BRowNE,
1789 edition, page 386) is as follows:
“SEPIA 1. Vagina subovato-truncata.
“THE INK or SCUTTLE-FISH
“This insect is frequent enough about Jamaica, but
most common on the north side of the island. It is
composed of a firm transparent sheath which in-
cludes the greatest part of an adherent but softer
gelatinous mass, furnished with a great number of
tentaculae of different sizes and forms. It is curious
to see how readily this creature discharges its ink on
the approach of danger, to hide itself in the coloured
fluid: but the juices discharged on such occasions,
are not only black and thereby sufficient to protect
the creature by giving a tincture to and thickening
the water about it; they are also bitter and clammy
which must probably render them either pernicious
to the gills, or hurtful to the eyes of all other fishes.”
Mr. Farr also wrote, “We do not know of any speci-
mens of Sepia having been found in Jamaica, however
cuttlebone has been found at Morant Point at the eastern
end of the Island. Mr. Lewis, the Director of the Institute
of Jamaica, tells me that his daughter used to collect it
there and bring it home for her budgerigars of which she
had a considerable number. So cuttle bone does not seem
to be rare on Jamaican Beaches.”
Dr. Ivan Goodbody of the Zoology Department, Uni-
versity of the West Indies, at Kingston, Jamaica, confirms
Mr. Farr’s account, writing to us: “I also have collected
cuttle bones at Morant Point washed up on the shore,
but in spite of doing some extensive Isaacs-Kidd trawling
and other fishing off the eastern end of the island none of
us have ever collected a living Sepia. As the cuttle bones
are very buoyant and resilient, they could be carried to
Jamaica from a very long way away. I would therefore
hesitate to suggest that the cuttle bones which we find
come from local animals.”
Vol. 12; No. 1
Still, even supposing that cuttlefish do not live in Carib-
bean waters, it is of interest that the sepions float to
Jamaica with some frequency, and have never been found
in Puerto Rico.
The case of cuttlebones on New Zealand beaches is
noteworthy. Sutrer (1913) listed 3 records of Sepia
apama Gray from there, each based on a single sepion
from the beach. Almost 50 years later, PowELL (1962, p.
71) noted that cuttlebones and shells of the chambered
Nautilus occasionally wash ashore on those islands, but he
did not think either cuttlefish or nautili lived there. His
list (op. cit.,p.126) of species of Sepia cuttlebones
found in New Zealand has only S. apama and S. plangon,
both known to live in Australia, some 2000 miles west-
ward. Cotton & Goprrey (1940) list 11 species of Sepia
from South Australia, including those two, and 29 others
found elsewhere around the continent. It is strange that
cuttlebones of only two of the many species in Australia
have floated to New Zealand. Perhaps it is a matter of
differences in living habits of the cuttlefish, or intrinsic
differences in the cuttlebones, or perhaps it is merely a
failure to record the cuttlebones from the shores of New
Zealand. ALLEN gives an interesting account of cuttlefish
behavior in Australian waters and the cuttlebones on the
beaches. She also notes that the sepion of Amplisepia
verreauxt ROCHEBRUNE, which is a senior synonym of
Sepia apama Gray, may reach a length of 45cm, or
18 inches.
There is one further bit of evidence that Sepia does
occur in the western Atlantic. Oxtverra (1940) named
a new subspecies from Brazil, Sepia officinalis jurajubai.
Her description is too meager to be of much use in differ-
entiating populations of this difficult species complex. In
the same year, Apam (1940) proposed that several nom-
inal species of larger cuttlefish of the eastern Atlantic are
merely subspecies of §. officinalis, and these can be in part
distinguished by their cuttlebones. Oxtverra did not de-
scribe or illustrate the cuttlebone of the specimen she
examined, but from her photographs it is evident that
she had a whole animal and that it is similar to S. off-
cinalis.
Could it be that Ottvemra’s specimen had migrated
across the Atlantic? If Sepia lives in the western Atlantic,
why is its presence so rarely reported? Perhaps it does live
here, but in situations which are little frequented by fisher-
men using suitable gear to catch them. ABEL (1916) made
a study of the habits and behavior of living cephalopods
for a better understanding of fossil species, but he merely
notes of Sepia that it lives near the bottom, over sand.
ABEL, ADAM, and other writers say that §. officinalis is
migratory, spending most of the year in deeper water,
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Page 91
but coming to shore during the breeding season, in the
warmer months. Tryon’s (1883, vol. 2. p. 40) account
is worth quoting:
“According to Verany, this animal (S. officinalis)
prefers rocky localities in the Mediterranean, where
it is fished by means of a dredge called a balancelle,
and is also taken at night with the trident. During the
month of March the fishermen use a living female
Cuttle fastened to a rope, or an imitation of one
formed of wooed, and made attractive to the male sex
by being ornamented with bits of glass; this latter
enveiglement is called by the Sicilians a fumedda,
and fishing with either of them is very productive
and amusing, especially on a moonlight night . ...
Their flesh is esteemed and abounds in the Italian
markets at all seasons of the year.”
Lane (1962, pp. 169ff) describes a fishing method of
baited traps used in the Mediterranean. These are made
of wicker baskets, attached along a weighted line. He also
repeats the account of fishing with female cuttlefish and
artificial lures.
From the above descriptions, we may suppose that if
Sepia lives near Galveston, it is likely to be around the
rocky reefs that occur beyond the 10-fathom zone, some
30 miles offshore. Those areas are not attractive to fisher-
men using shrimp trawls, because of the danger of
re“
ee
Figure 1
Ventral view of the shell of Sepia mestus Gray.
After Hoyze, (1886, p. 123)
1-Last loculus - 2-Chitinous margin of outer cone
3 - Striated area (margins of earlier loculi) ~ 4-Limb of inner cone
5 - Inner cone (a thin filet of calcareous matter) - 6- Spine
Page 92
THE VELIGER
Vol. 12; No. 1
Figure 2
Ventral view of a cuttle bone found at Galveston. 70 mm long
tearing the nets. If the population of Sepia were small,
the individuals alert but shy, they might not be easily
taken.
The cuttlebones of Galveston (Figures 2 and 3) seem
to be all of one species; there is nothing very different
from the specimens of Sepia officinalis ossicles sold in the
pet shops, nor from descriptions and illustrations of that
species in the literature. Figure 1 indicates the terms ap-
plied to parts of a cuttlebone. Figures 2 and 3 were drawn
by the junior author from the beach specimens of Galves-
ton.
Figure 3
Dorsal view of same cuttlebone shown in Figure 2
The outline is elongate oval, with the sides almost paral-
lel throughout most of their length, and slightly sinuate
about a fourth of the length from the hind end. The
locular margins were too worn to note the pattern of
their edges or the locular index (length of last loculus
as percent of total length). The dorsal surface is covered
by a thin sheet of shiny, transparent cuticle over its hind
end, and the rest is uniformly, faintly roughened by short,
transverse, closely spaced rugae. There is a shallow furrow,
poorly defined, on each side of the midline, with a low
medial ridge between them. In side view, the cuttlebone
Vol. 12; No. 1
is almost flat. The spine is short and acute. The chevron
markings on the ventral surface of the one drawn seem
to be the bill print of some shore bird.
Measurements (in millimeters)
Length Width
70 22 (Figures 2, 3)
82 26
88 30
75 25
75 26
That there is much yet to be discovered concerning
the mollusks, and particularly the cephalopods, of the
Gulf of Mexico can scarcely be denied. We may further
note that Voss (1956), in monographing the cephalo-
pods of the Gulf, merely suspected the presence of
Argonauta here, and had no specimens from the area.
Several specimens with animal and shell were found in the
stomach of the dolphin, Coryphaena, in the summer of
1966 by Mr. Jim Dailey, while fishing south of Freeport,
Texas. To our knowledge, Argonauta has never been found
alive in this area. Beach shells are much rarer than the
literature indicates. At least, we have never found any in
Louisiana or Texas. But Coryphaena knows where the
paper nautilus lives, and how to capture it. WARMKE
(1961) also reports the paper nautilus from a_fish’s
stomach in Puerto Rico.
Interesting in this respect is the case of the giant squids
of the family Architeuthidae, monsters of 4m _ body
length and more. These are known only from disabled
specimens floating on the surface, or cast ashore, or dis-
gorged by captured whales. Lance (1962) has a vivid
account of battles between these squid and whales, noting
that the squid may be the protagonist (see also VERRILL,
1882). Although giant squid are said to be cosmopolitan,
the dearth of records of juveniles captured by trawling
is striking, particularly if they are, as noted by Voss
(1956, p. 138): “in all probability very poor or weak
swimmers and not at all adapted for catching active prey.
Indeed, they should fall easy victims to the sperm whale,
their natural enemy.”
ADDENDUM
After this paper was written, Mr. William Wardle found
two cuttlebones on the east end of Galveston Island, on
21 December 1968. These are about equal in size, but
they are larger than the ones found nearly a year before.
The larger one is 135mm long, 43mm wide, and 19mm
THE VELIGER
Page 93
thick. They are of the same species as those found earlier,
and in even more perfect condition. There is no evidence
of epibiota on them. A break in the inner cone of one shows
a transparent material resembling cartilage filling its
interior. The lower surface of both specimens was badly
scarred and pitted by chevron markings, again suggesting
these had been pecked by sea birds. The lower surface is
thickest about one-fourth the length from the hind end,
from which point it slopes forward in a gentle convex
arch, and backward in a more pronouncedly concave
excavation. The striated area begins at the thickest point
of the sepion, and thus the locular index is about 75%
of the length.
Besides being larger, the Wardle specimens show two
important anatomical differences from the smaller ones:
(1) The dorsal surface has a thin, smooth cuticle along
the posterior half of each margin, leaving a broad V-shaped
area in the middle free of it. The apex of the V is nearly
at the hind tip of the shell. Contrast the shallow V of the
smaller specimen drawn in Figure 3, in which the dorsal
cuticle is limited to the hind fourth of the shell.
(2) There is no projecting spine, and the chitinous mar-
gin of the outer cone seems to extend a little farther
behind the calcareous part of the shell than in the smaller
specimens. These differences probably represent changes
during growth, rather than distinct species.
The sculpture of the dorsal area is of small bosses and
short vermiculate rugae, rather irregularly oriented and
closely spaced. The Wardle specimens were dropped into
alcohol when found. This method of preservation allows
details to be seen much more clearly than in the smaller
specimens, which were dried.
LITERATURE CITED
ABEL, O.
1916. Paldobiologie der Cephalopoden. Gustav Fischer,
Jena; 281 pp.
ApaM, W.
1940. Les races de la seiche commune (Sepia officinalis Linné).
Bull. Soc. Zool. France 65: 125 - 131
ALLAN, Joyce K.
1950. Australian shells.
Browne, PATRICK
Georgian House, Melbourne; 470 pp.
1756. The civil and natural history of Jamaica: viii +503
pp.; 49 plts. (second ed. 1789: viii+490 pp.)
Bruun, ANTON, Jr.
1943. The biology of Spirula spirula (L.). Dana Reprt.
No. 24: 1 - 44
Cotton, BERNARD CHARLES & FRANK K. GopFREY
1940. The Mollusca of South Australia. Part II - Scaphopoda,
Cephalopoda, Aplacophora and Crepipoda. Frank Trigg,
Printer, Adelaide. pp. 315 - 600
Page 94
ErpMAN, DonaLp S.
1957. | Vagabond cuttlebones.
Forses, Epwarp & SYLVANUS HANLEY
1848-1853. A history of British Mollusca and their shells. Van
Vorst, London; 4 vols.; illustr.
Hoy.e, WiLLiAmM Evans
1886. Report on the Cephalopoda collected by H.M.S.
Challenger during the years 1873 - 76. Rept. Sci. Res.
Voy. Challenger 1873-76. Zool. 16 (44): 1-245; 33 plts.;
10 figs. in text
Lane, Frank W.
1960. Kingdom of the octopus.
(1962 ed.), 287 pp.
O.iverra, HELENA Paks DE
1940. Sobre dois cefalopodos da Guanabana.
Agric. (Rio de Janeiro, Brazil) 10: 29 - 35
Orsicny, Atcipe DESSALINES D’
(1841) -1853. Mollusques, in Ramon de la Sagra, Histoire phy-
sique, politique et naturelle de l’ile de Cuba. Paris (Arthus
Bertrand) : 2 vols., atlas of 28 plts.
PowELL, ARTHUR WILLIAM BADEN
1962. Shells of New Zealand. Whitcombe & Tombs, Auck-
land, 4‘ ed., pp. 1 - 203; pits. 1-36; text figs.
Nautilus 70 (3): 106 - 107
Pyramid Publ. New York
Bol, Minist.
THE VELIGER
Vol. 12; No. 1
Smrrn, Epcar ALBERT
1916. | On the shells of the South African species of the Sepi-
idae. Proc. Malacol. Soc. London 12: 20 - 26; plt. 2
Suter, HENRY
1913. Manual of New Zealand Mollusca. MacKay,
Wellington, New Zealand, 1 - 1129; atlas of plates (1915)
Tryon, GEORGE WasHINGTON, Jr.
1883. Structural and systematic conchology: an introduction
to the study of the Mollusca 2. Philadelphia. pp. 1 - 430;
pits. 23 - 91
VERRILL, ADDISON E.
1882. Report on the cephalopods of the northeastern coast of
America. Ann. Reprt. U.S.Comm. Fish & Fisheries for
1879: 1 - 244; 46 plts.
Voss, GrcBerT L.
1956. A review of the cephalopods of the Gulf of Mexico.
Bull. Mar. Sci. Gulf and Carib. 6 (2): 85-178; 18 figs.
WarRMKE, GERMAINE L,
1961. Brown paper nautilus recorded from fish stomach in
Puerto Rico. Caribb. Journ. Sci. 1 (4): 142 [not seen]
WINcKworTH, RONALD
1936. Marine Mollusca from South India and Ceylon. IV. A
new Indian Sepia.
pit. 4
Proc. Malacol. Soc. London 22: 16 - 17;
Vol. 12; No. 1
THE VELIGER
Page 95
A Revision of the Eastern Pacific Ovulidae
BY
CRAWFORD N. CATE
12719 San Vicente Boulevard, Los Angeles, California 90049
(Plates 7 to 10; 3 Maps)
INTRODUCTION
REcENTLY I was ASKED to identify some Simnia from the
Gulf of California. During the identification of these
species, much confusion was noted and it seemed that
some attempt to clarify the nomenclatural status of them
would be desirable. In examining the shells in private col-
lections and West Coast Museums I observed that many
remained unidentified while others were incorrectly iden-
tified. In consulting the literature it soon became obvious
that little work had been done with these shells in recent
years, and what had been accomplished earlier gave no
clear indication of the essential differences existing be-
tween the species. An attempt will be made here to
show how the shells of this group of species are distinctly
separable from one another; the names of those that do
not meet this requirement will be relegated to synonymy.
A part of the eastern Pacific Ovulidae comprise a
group of 10 species assigned to 3 genera: Jenneria,
Simnia, and Cyphoma. An attempt will be made here to
identify the species by means of specific morphological
and color characteristics, employing the nomenclature
proposed by ScuILpErR (1968) ; the species are illustrated
and a limited locality list is provided to show the ranges
of the species within the prescribed area and the manner
in which some of these ranges overlap.
DISCUSSION
In the two west American species, Simnia loebbeckeana
(Weinkaurr, 1881) and S. catalinensis (BERRY, 1916),
the shells are, with few variations, ivory colored; in
Cyphoma emarginata (SowErRBy', 1830) the shell is con-
sistently off-white in color. In the other 7 species, however,
color is a basis for confusion, particularly where the shell
assumes the color of the host gorgonian upon which the
species lives; these colors may vary from white to an
intense deep purple-brown. There is one exception among
them, however; in S. aequalis (SoweRBY", 1832), the yel-
low color on white shells, and yellow-orange on the tinted
specimens, constitutes a major point in the identification
of the species.
Some of the animals in Simnia, although having appar-
ently identical shells, seem to vary in the color of the soft
parts. It is generally assumed that simniids often take on
the color of the gorgonians on which they live.
In the sub-group Cyproglobinini ScuiLprEr, 1932, there
is one accepted genus: Jenneria JoUSSEAUME, 1884. In
the sub-group Simniini ScHiLpER, 1927, there are two
accepted genera: Simnia Risso, 1826, and Cyphoma
Ropinec, 1798. Neosimnia Fiscuer, 1884, has been rele-
gated to synonymy with Simnia (ScuiLpeEr, 1968, p. 271)
and therefore will not be used in this report.
GENERIC KEY
Cyproglobinini: Shells cypraea-form; surface finely stri-
ate; dorsum nodulate Jenneria
Simniini: Shells elongate; terminals usually short (shorter
than in Volva Ropinc, 1798) ; anterior aperture less con-
stricted, or not at all; transverse carina on dorsum absent.
Outer lip narrow or sharply edged, inflected or reflected;
with or without fossula Simnia
Shell long, wide; terminals broad, rounded or subsquare;
shell thickly, solidly constructed; central dorsal carina
prominently elevated; a funicular, carinal twist on the
adapical surface of the columella may be present
Cyphoma
(Jenneria and Cyphoma include but a single species each
in the area covered by this report.)
KEY to THE SPECIES or Simnia
Smooth, glossy, without dorsal sculpture ..... loebbeckeana
White, sharply upraised columellar carina along entire
length of shell rufa
Page 96
THE VELIGER
Vol. 12; No. 1
Dorsal sculpture of upraised transverse ridges covering
entire mipperishellsuntace er ete eee eter avena
Dorsal surface only partially sculptured with transverse
ridges emanating from both terminals .......... catalinensis
Transverse ridges emanate restrictedly from each termin-
al; yellow terminals on white shells, orange terminals
on colored shells; columellar carina weakly, finely
formed ie ceine tact Meee et eh Ae aequalis aequalis
Transverse ridges emanating from each terminal; yellow
to orange terminal tips; pronounced, upraised, white
carina on forward half of columella ..... aequalis vidleri
Transverse ridges emanating away from each terminal,
and yellow terminal tips; shell broader, with a wider
curving carina on the forward half of the columella
oH ten Uren eee eae ieee bellamaris
LOCALITY INDEX
Many of the localities listed here were obtained from
handwritten labels in various collections. Every effort has
been made to verify the spellings, but in some instances
certain place-names were not found on any map or atlas
available to me; it is regretted if any discrepancies occur.
1. Aguachale, NE Baja California del Norte; approx.
24 miles S of San Felipe
2. ibid.; in 5 feet of water
3. ibid.; 2 miles S of —
4. Albemarle Island (= Isabella Island), Galapagos
Islands, Ecuador
5. Avalon, Catalina Island, S. California; in 40 fathoms
6. Bahia de Adair, Sonora, West Mexico; approx. 20
miles N of Punta Penasco
7. Baja Isla Grande (= Tiburon Island), Gulf of Cali-
fornia
8. Bird Island, Catalina Island, S. California; in 35 to
40 feet of water
9. Calito de Campos, Michoacan, Mexico; N of Aca-
pulco
10. Calito Mero, Peru
11. Carpinteria, California; deep water
12. Catalina Island, California; approx. 22 miles W of
San Pedro
13. ibid.; at the Isthmus in 35 feet of water
14. Cerralvo Island, SE Baja California; just E of La Paz
15. Concepcion Bay, E Baja California del Sur; just S of
Mulege
16. Corona del Mar, California ; just S of Newport Beach
17. Creston Island, Mazatlan, Sinaloa, Mexico; in 25 feet
of water, on gorgonians
18. Deer Island, Guaymas, Sonora, Mexico; in deep water
. Dominica, West Indies; Monte Christi Beach
. Estero Soldado, Guaymas, Sonora, Mexico
. Galapagos Islands, Ecuador
. Guaymas, Sonora, Mexico
. Key Largo, Florida Keys, Florida
. La Abreojos, W Baja California del Sur, approx.
26°40’ N Lat.; 113°35’ W Long.
. Laguna Beach, Orange County, California; in 40 to
50 feet of water
. Las Gaviotas, Mazatlan, Sinaloa, Mexico
. Magdalena Bay, SW Baja California del Sur
. Manzanillo, Colima, Mexico
. ibid.; El Dorado Bay
. Mazatlan, Sinaloa, Mexico
. Mission Bay, San Diego, California
. Monterey Bay, Monterey, California
. Morro Bay, Pismo Beach, California; in 80 feet of
water
. Newport Beach, California
. ibid; off breakwater in 20 - 25 feet of water
. Ocean Park, California, adjacent to Venice: 5 miles
W in 300 feet of water (J. L. Baxter)
. Pajaros, Mazatlan, Sinaloa, Mexico
. Panama Bay, Pacific Panama, W Central America
. Panama City, Panama; in deep water
. Perlas Islands, Panama Bay
. Playa del Rey, California; in 180 feet of water
. Punta Diggs (Punta Estrella), E Baja California del
Norte; approx. 6 miles S of San Felipe
. Point Fermin, San Pedro, California
. Puertecitos, E Baja California del Norte; approx. 75
miles S of San Felipe
_ Puerto Escondido, E Baja Califomia del Sur; 11 miles
S of Loreto
. Pulmo Reef, SE Baja California del Sur; approx. 70
miles S of La Paz
. Punta Cameron, Mazatlan, Sinaloa, Mexico
. Punta Penasco (Cholla Bay), Sonora, Mexico
. Punta San Ignacio, Punta Penasco, Sonora, Mexico
. Redondo Beach, California; in 100 feet of water
(R. Mistrell)
. Santa Barbara, California
. Santa Cruz, Nayarit, Mexico; 15 miles S of San Blas
. San Carlos Bay, Guaymas, Sonora, Mexico; dredged
in 17 fathoms
. San Diego Bay, San Diego, California; entrance
. San Felipe, E. Baja California del Norte
. San Francisco Bay, Guaymas, Sonora, Mexico
. ibid.; dredged from 16 fathoms
. San Onofre, California; approx. 18 miles N of Ocean-
side (C. C. Finlay)
. San Pedro, California (Mrs. L. C. Oldroyd)
THE VELIGER, Vol. 12, No. 1 [C. N.CateE] Plate 7
Figure 1 Figure 2
Simnia rufa (SowERBy 24, 1832) Simnia rufa var. inflexa (SowERBy 24, 1832)
Pulmo Reef xs 3 Estero Soldado x34
Figure 3 Figure 4
Simnia aequalis aequalis (SowERBy 2"4, 1832) Simnia aequalis vidleri (SowERBY 34, 1881)
Point Diggs xX 2% Morro Bay 25
Figure 5 Figure 6
Simnia avena (Sowersy 24, 1832) Simnia bellamaris (BERRY, 1946)
Creston Island xX 5 San Diego Bay X 3
photographs by R. PoorMaN
Vol. 12; No. I
THE VELIGER
Page 97
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60. ibid.; off breakwater
61. Santa Monica, California; 44 miles W of (J. L. Baxter)
62. Santa Rosalia, E Baja California del Sur
63. Saladita Bay, Guaymas, Sonora, Mexico
64. Tabago Island, Honda Bay in Panama Bay; 12 miles
SW of Panama
65. Venado Beach, Panama, Central America
66. Venado Island, Panama, Central America
Jenneria pustulata (LicutFoot, 1786)
Cat. Portland Mus., London 1786 (2230) :
106 — (not of SoLaNnpER, 1786; nor
Lamarck, 1810)
(Plate 8, Figure 9)
Type Locality here designated: Cholla Bay, Punta
Pefiasco, Sonora, Mexico (30°40’ N Lat., 113°20’
W Long.)
(emend. China: Lister error; LicHTFoot, 1786)
This is a fairly common species in some localities; it ranges
from the upper Gulf of California south to Ecuador
(KEEN, 1958). It has been demonstrated that the species
is most closely related to the Ovulidae (see D’Asaro,
1969), and it is therefore included in this report. The
shell shape of Jenneria pustulata is that of a cypraeid;
the apertural teeth traverse both base and outer lip as
sharply elevated white ridges; the pale grey dorsum is
divided by a mantle line, and the overall dorsal surface is
thickly overlaid with bright orange pustules, which are
encircled by a brown ring — there are two large brown
spots superimposed over the area inward from each ter-
minal. The shells average about 20 by 12.5 by 8.2 mm in
length, width, and height, respectively. Those specimens
that I have seen live in the sand, digging into it with the
the receding tide, at Cholla Bay.
Localities: 2 3 22 26 29 38 44 45 52 65
Simnia rufa (SowERBY", 1832)
Conch. Illust., London: fig. 58
(Plate 7, Figures 1, 2; Plate 9, Figures 13, 14)
Syn.: Ovulum inflecum SowErBy", 1832
Conch. Illust., London, fig. 60
Ovula californica Reeve, 1865 (Sowersy MS)
Conch. Icon., 15 Ovulum, fig. 50
Ovula neglecta Reeve, 1865
Conch. Icon., 15 Ovulum, fig. 62
This is a fairly common, distinctive species. It differs
from other members of the genus by having a generally
narrower shell; by being rectangularly elongate; by having
Page 98 THE VELIGER Vol. 12; No. 1
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THE VELIGER, Vol. 12, No. 1
Figure 7
Simnia loebbeckeana (WEINKAUFF, 1881)
Magdalena Bay Xx 2
Jenneria pustulata (Licutroort, 1786)
Puertecitos X 2%
Figure 11
Cyphoma intermedia (SowERBy ***t, 1828)
Dominica x 12
photographs by R. PoorMAN
[C. N.CateE] Plate 8
Figure 8
Simnia catalinensis (BERRY, 1916)
Avalon DS Ore
Figure 10
Cyphoma emarginata (Sowerby ', 1830)
Puertecitos
Figure 12
Cyphoma gibbosa (LINNAEUS, 1758)
Key Largo xX 2
Vol. 12; No. 1 THE VELIGER Page 99
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Page 100
a smooth dorsum without any important sculpturing
(sometimes exceedingly faint growth lines are visible at
the terminals) ; but most importantly it differs by having
an upraised carinal ridge the full length of the columella.
The shell may be either white or of varying shades from
purple-brown to a striking deep purple; the shell mar-
gins may have an orange cast at times.
The shell varies in length up to about 24mm; the
animal lives on gorgonians and sea-fans, and will likely
acquire the color of the host colony on which it lives.
The species occurs abundantly at Guaymas and the off
shore islands and reefs of southeastern Baja California.
Localities: 6 7 14 15 18 20 22 24 28 30 37 38 44
46 49 53 59 62 63 64
Simnia avena (SOwERBY", 1832)
Conch. Illust., London, fig. 59
(Plate 7, Figure 5)
Syn.: Ovula livida ReEve, 1865
Conch. Icon., 15 Ovulum, fig. 63
This species seems to be uncommon. It is easy to identify
because of the general appearance of the shell; it is a
small form, rarely measuring more than 14 mm in length.
It differs from other Simnia species by being broad for its
length; and, most importantly, the shell is ribbed over
the dorsum, transversely from one terminal to the other.
These sharply upraised ridges are the most convenient
means of identification.
As it is presently known, the species seems to be restric-
ted to the west coast of Mexico and Central America to
Panama, although specimens have been reported from
Calita Mero, Peru; also the species is well established at
the Galapagos Islands.
Localities: 10 17 20 21 30 37 38 56
. Simnia aequalis aequalis (SOWERBY ", 1832)
Conch. Illust., London, fig. 61
(Plate 7, Figure 3; Plate 9, Figures 15, 17)
Syn.: Ovula variabilis REEvE, 1865 (C. B. ApAMs MS)
Conch. Icon., 15 Ovulum, fig. 60
Neosimnia quaylei Lowe, 1935
Trans. San Diego Soc. Nat. Hist. 8 (6):
74 Din By IHR,
Neosimnia vidlert tyrianthina Berry, 1960
Leafl. Malac. 1 (19): 118 (31 Dec.)
This is possibly the most common Simnia species occurring
in the Panamic region, with the center of abundance along
the shores of northeast Baja California, and appearing
to occur to a limited extent in the northern half of the
THE VELIGER
Vol. 12; No. 1
Gulf of California. Although the species seems to have
many synonyms, it appears to possess a distinctive color
and form (this appears to be the only simniid in which
color can be used as a criterion for identification). The
shell varies in length up to about 19mm; the animal
lives on variously colored gorgonians.
This species differs from its congeners by having yellow
terminal tips on white shells, the color blending into
dull orange on shells of rose and lavender-brown; it dif-
fers further by having transverse ribbing on the dorsal
surface radiating away from the terminal openings; and,
unlike its subspecies, it has a very weakly formed carinal
ridge on the forward half of the columella. This combi-
nation of shell characters is missing from all other eastern
Pacific Simnia, but is present in all of the mentioned
synonymic species.
Localities: 2 3 6 20 22 42 44 45 53 55 57
Simnia aequalis vidleri (SowERBY™, 1881)
Proc. Malacol. Soc. London 1881: 683; plt.
56, fig. 1
(Plate 7, Figure 4; Plate 9, Figure 16)
Syn.: Ovula subrostrata (SowERByY", 1832)
Proc. Zool. Soc. London 1848: 136
This subspecies is fairly common below the low tide mark,
and is found living on gorgonians and related organisms.
The subspecies is geographically separated from the nom-
inate species, which lives in the northern reaches of
the Gulf of California. Although having the color and
morphological characters of the species in the strict sense,
it is separated by possessing a heavily-formed, white,
sharply upraised carinal ridge that traverses the forward
half of the columella. It is this feature, along with its
West American locality, that separates it from all other
Simnia forms. These shells also seem to grow longer and
broader, varying in length up to 25 mm. Szmnia aequalis
vidleri appears not to occur in the Gulf of California,
but to be limited to American coastal waters, including
the offshore islands, from Monterey Bay, California, to
the west coast of Central America, Panama, and the
Galapagos Islands.
Localities: 4 21 22 24 30 32 33 35 37 38 40 50
Simnia bellamaris (Berry, 1946)
Journ. Conch., Manchester 22 (8): 190;
fig. 1
(Plate 7, Figure 6; Plate 10, Figure 21)
This species is rare and seems presently to be restricted to
the bays adjacent to San Diego, California. It is broader
THE VELIGcER, Vol. 12, No. 1 [C. N.Cate] Plate 9
oa pecho a, Sour i
Californienm_ Wo. o [ - Af Cexa, Sow -
Figure 13 ay a
Simnia rufa (SowErRBy 2"4, 1832) Simnia rufa (SowErBy 24, 1832)
Syntypes X 2 inflexa syntypes x 2
Sam AAW ot
15a Simnia aequalis aequalis (SowERBY 24, 1832) Figure 16
15b Simnia avena (Sowersy 74, 1832) Simnia aequalis vidleri (SowERBy 3°, 1881)
BM(NH): No. 81.5.20.31 | Monterey X 2
cnhractratinm.. Samnerk
igure 18a Simnia aequalis aequalis (SowERBY 2™4, 1832)
Simnia aequalis aequalis (SowERBy 7", 1832) 18b,c,d Simnia avena (Sowersy 2"4, 1832)
(rufa-subrostratum)
photographs by A. Myra KEENn
iy
THE VELIGER, Vol. 12, No. 1 [C. N.CarE] Plate 10
ie ee NS AS SN tee Gee Ge ee
Figure 19 Figure 20
Simnia aequalis aequalis (SowERBy 2"4, 1832)
Simnia aequalis aequalis (SowERBy 2°4, 1832)
(tyrianthina Berry ) Holotype 16mm
(quaylei LowE) Type SDNHM No. 602 xX 3
No...7846..270 7
Neojimnia bellé sosimr o cotal
iwalon .
4
Figure 22
Simnia catalinensis (BERRY, 1916)
Type (24 mm)
Figure 21
Simnia bellamaris (BERRY, 1946)
a - Holotype (26 mm) b - Paratype (22 mm)
photographs by James H. McLEAN
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Vol. 12; No. 1
with a wider aperture than the other eastern Pacific
Simnia, being particularly wide for the length of the shell;
the average measurements of shells examined were 18 mm
long and 13 mm wide. The species possesses many of the
morphological features of S. aequalis vidleri (see above),
but seems to differ from it mostly in the dimensions of
the shell and by a more heavily formed columellar carina.
Otherwise it very closely approaches S. aequalis vidlert.
Since S. bellamaris might well be a localized morph-
form of this species, it is listed here provisionally.
Localities: 31 54
Simnia loebbeckeana (WEINKAUFF, 1881)
Syst. Conch. Cab., Mart. Chemn. ed., 5/3:
197; plt. 50, figs. 6, 7
(Plate 8, Figure 7)
Syn.: Ovulum fornicarium Keep, 1887 (not of SowER-
By ', 1828) ; West Coast Shells 1887: 61; fig.
47
Ovula barbarensis Dati, 1892
Proc. U.S. Nat. Mus., 15: 206; plt. 21, fig. 1
This is a fairly common species, ranging along the West
American coast from Magdalena Bay, Baja California,
north to Monterey Bay, California. The shell is almost
always ivory colored, although some specimens examined
showed a faint pink blush of color. The average shell size
is about 19 mm, but an occasional specimen may meas-
ure 24 mm in length (Calif. Acad. Sci. Coll.).
The species is distinct, with a morphology that seems
distantly related to a species occurring in deep water off
Catalina Island, Simnia catalinensis (Berry, 1916). It
differs from this species, however, by being generally smal-
ler and narrower; and, most importantly, by having no
dorsal sculpturing, the shell being smooth and glossy
throughout.
Localities: 11 16 27 32 36 41 43 51 58 59 61
Simnia catalinensis (BERRY, 1916)
Nautilus 30 (2): 21; plt. 5, fig. 3
(Plate 8, Figure 8; Plate 10, Figure 22)
This deep water species is not common, and presently
seems to be restricted to the offshore environs of Catalina
Island, California. The shells are large, lightweight in
construction and bulbously broad. Average shell length is
25 mm and the breadth is 14.5 mm; the shell color is pale
ivory-white; it differs most importantly from Simnia loeb-
beckeana by having transverse rib sculpturing on the dor-
sum at either end-quarter of the shell; it is also a larger
THE VELIGER
Page 101
simniid species. Simnia loebbeckeana, the previously dis-
cussed species, has a smooth dorsal surface without trans-
verse ribbing.
Localities: 5 8 12 13
Cyphoma emarginata (SowerBy', 1830)
Spec. Conch. 1: Ovulum 1: 7; plt. 54, figs.
54, 55
(Plate 8, Figure 10)
Syn.: Cyphoma marginata CHENU, 1859 (used in error)
Man. de Conchyl. 1: 273; figs. 1789, 1793
This is a relatively common species in certain localities.
There may be some relationship between this form and the
West Indian Cyphoma gibbosa (Linnaeus, 1758), Plate
8, Figure 12, especially the form called C. intermedia
(Sowersy', 1828), Plate 8, Figure 11. This latter species
may be only an uncalloused intermediate to the heavily
margined form mentioned above.
The Panamic shells are off-white to whitish beige, with
a pale beige blush at the terminal openings; adapical
funicular cord gives the posterior terminal beak a peculiar
twist to the right; however, the most important shell char-
acter is the transverse dorsal ridge, which identifies the
species generically. Cyphoma emarginata seems to be
the only species in this genus occurring in the eastern
Pacific region. Shells vary up to 28 by 16 by 12.5 mm in
length, width, and height, respectively.
Localities: 3 9 20 44 47 — Cyphoma intermedia: 19
Cyphoma gibbosa: 23
ACKNOWLEDGMENTS
In the preparation of this paper I enjoyed the help and
assistance of many people and I wish to thank Dr. Myra
Keen, Stanford University; Dr. Leo Hertlein, California
Academy of Sciences; Dr. James McLean, Los Angeles
County Museum of Natural History; Dr. George Radwin,
San Diego Museum of Natural History; Dr. Takeo Susuki,
University of California at Los Angeles; and Dr. Eugene
Coan, Stanford University: for the use of the collec-
tions, for their counsel and for help with access to obscure
literature. To several private collectors go my thanks for
the use of their collections from the Gulf of California;
these especially include Twila Bratcher, Roy and Forrest
Poorman, Helen DuShane, and Gale Sphon.
I wish to extend special thanks to Dr. Myra Keen for
the use of photographs taken by her of the simniid types
in the British Museum (Natural History) (Plate 9, Fig-
ures 13 to 19) ; to Dr. James McLean for additional type
Page 102
THE VELIGER
photographs from his institution (Plate 10, Figures 19 to
22); to Roy Poorman for photographing the specimens
illustrated in Plate 7 and 8 (Figures 1 to 12) ; to Dr. Takeo
Susuki for the special processing of the photographs; and
to Mrs. Emily Reid for the excellently drawn maps. To
Jean Cate goes a very special thanks for her interest in
my work, and the assistance she has given in many ways,
all of which make for the success of this paper.
LITERATURE CITED
D’Asaro, Cuarzes N.
1969. The egg capsules of Jenneria pustulata (LicHTFOOT,
1786), with notes on spawning in the laboratory. The
Veliger 11 (3) : 182 - 184; 1 text fig.; 1 table (1 January 1969)
TREDALE, Tom
1935. Australian cowries, pt. I. Austral. Zool., 8 (2): 96 to
135; plts. 8-9
KEEN, A. MyRA -
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi + 624 pp.; illus.
Stanford, Calif. (Stanford Univ. Press)
LicHTFooT, JoHN
1786. A catalogue of the Portland Museum, lately the property
of the Duchess Dowager of Portland, deceased ... . London.
pp. i- viii+ 1-194
ScHILper, FRANz ALFRED
1968. The generic classification of cowries. The Veliger
10 (3) : 264-273 (1 January 1968)
Sowersy, GEorGE BRETTINGHAM (2%? of name)
1832. Conchological illustrations. Figures 1 - 61
Vol. 12; No. 1
—
Vol. 12; No. 1
THE VELIGER
Page 103
The Eastern Pacific Cowries
CRAWFORD N. CATE
12719 San Vicente Boulevard, Los Angeles, California 90049
(Plates 11 to 15; 3 Maps)
INTRODUCTION
THE PURPOSE OF THIS PAPER is to provide a checklist of
the cowrie species known from the Eastern Pacific Region
of the Americas, including such islands and island groups
as the Galapagos, Clipperton, Cocos, Revillagigedos and
Guadalupe. All of these offshore localities share a similar
molluscan fauna. It seems pertinent, therefore, to mention
a few facts regarding these islands.
First in geographical importance are the Galapagos Is-
lands, which provide a home for many cowrie species in-
cluding both eastern and western forms. The archipelago
consists of 10 major islands, the largest of which is Isa-
bella (Albemarle). Other more important islands, with
their earlier names are: Santa Cruz (Indefatigable), San
Cristobal (Chatham), San Salvador (James), Fernan-
dina (Narborough), Santa Maria (Charles), and Es-
pafiola (Hood). The collecting localities at these islands
are, for the most part, horizontal strata, lava outcroppings,
boulder strewn extensions of black basaltic lava with at-
tached algae, and a minimum of sandy beaches.
Clipperton Island appears to be the crossroads for the
eastern and western cowries, as here the two faunas come
together. This former atoll is located on the western peri-
meter of the Eastern Pacific cowrie region and seems to
provide a bridge that has enabled some of the more
western species to migrate to the American mainland.
Clipperton Island is roughly 2 square miles of irregular
atoll land area; it is uninhabited and is about 670 miles
southwest of Mexico. With a diameter of about 24 miles,
it encloses a rocky projection of about 85 feet in height,
the highest point on the atoll. The shore area appears to
be ideal for cypraeids with its sandy beaches, some of
which are strewn with rock and coral rubble, and the
adjacent coral and algae reef-flats.
Luria isabellamexicana (STEARNS, 1893) seems to be
the native cowrie species, considering the east and west
extremes of its range. It appears to reproduce in greatest
abundance at Clipperton Island, whence its range extends
eastward to the west Mexican-Central American inter-
tidal waters. Westward the species emerges again in Ha-
waiian subtidal waters as L. isabella controversa (Gray,
1824). This range determination becomes more valid
when one considers that L. isabella atriceps SCHILDER &
ScHILper, 1938, a clearly distinguishable subspecies, can
be observed to commence its West-Central Pacific range
in the Hawaiian Islands, a fact that should not have been
ignored in Cate (1965). A study of a series of these sub-
species shows little or no intergrading between them, even
though they share overlapping habitats.
Cocos Island, Costa Rica, is an uninhabited island of
approximately 18 square miles, situated southwesterly
from the mainland. Not much field work has been under-
taken at this presently inaccessible locality, and, therefore
there is not much known of its cowrie fauna.
The Revillagigedos Islands lie about 420 miles west of
the Mexican mainland and, nearly 240 miles south of
Cape San Lucas, Baja California del Sur. These islands
belong to Mexico, and consist mainly of Clarion and So-
corro Islands; Roca Partida Island is the westernmost of
this group (see HERTLEIN &« Hanna, 1930). Some field
work has been accomplished at these islands, resulting in
a somewhat better knowledge of the cowries there. How-
ever, isolation and inaccessibility still limit the field work
in this area.
The Tres Marias Islands, Mexico, on the other hand,
are only about 50 miles off the mainland coast of the
State of Nayarit, and some 200 miles southeast of Cape
San Lucas (Strone « Hanna, 1930). Several collecting
expeditions to these islands have fairly well explored the
fauna, so that we have an adequately complete knowledge
of the cowrie species occurring in these islands.
The Pacific coast of Central America represents the
connecting link, or land bridge, between North and South
America; it extends officially from the southern boundary
of Mexico to the northwest boundary of Colombia. This
coastline includes numerous bays, islets, clif-bound shores,
and rocky projections into the sea; coral tables at the
Page 104
ocean surface, coral reefs, and extensive sandy beaches.
Mangroves line areas of quiet backwaters; to all of this
is added a tropical water temperature. There are at least
5 of the Eastern Pacific cowrie species living within this
warm-water range. The northern coast of Pacific South
America provides for at least 2 more, with the remainder
confined to the Gulf of California-West America, and the
listed islands.
The Gulf of California, an arm of the Pacific Ocean,
extends northwest into the mainland of North America.
It is bounded on the East by mainland Mexico, and on the
West by the peninsula of Baja California. It is interesting
to note that most of the cowrie species in this province
thrive within the confines of the Gulf as well as along
the open shoreline. The one exception is Zonaria annettae
annettae (Dati, 1909), which appears to be completely
restricted to the waters of the Gulf.
The west coast of North America from Monterey, Cali-
fornia in the North to the proximity of Cedros Island,
West Baja California, in the South, provides another de-
limited range for cowries. Perhaps the most notable fac-
tor in the living conditions in this area are the cold water
upweilings, a condition not generally found in the other
areas included in this discussion. Zonaria (Neobernaya)
spadicea (Swainson, 1823) may be collected at nearly
any point in this range that provides suitable ecological
conditions for cowries; this includes all of the islands such
as the southern Guadalupe Island.
Guadalupe Island, a Mexican island possession, is situ-
ated about 275 miles west of El Rosario, Baja California
del Norte, and about 185 miles southwest of San Diego,
California. The island is surrounded by very deep water,
up to 2000 fathoms; it is of volcanic origin. It is said to
possess much the same marine ecological conditions as
those observed on the California coast and Californian
islands (Stronc & Hanna, 1930; CHace, 1958).
No provision is made here to include a report on the
possibility of cypraeids occurring at the remote, southern
Chilean islands of San Felix (20°20’S Lat.; 80°10’W
Long.) ; San Ambrosio (26°40’S Lat;80°00’W Long.) ;
and Juan Fernandez (33°30’S Lat.; 79°00’W Long.).
Special interest attaches to the discovery of two cowrie
species from the western Pacific on the beaches of Baja
California del Sur by Mrs. Helen DuShane, Whittier, Cali-
fornia. On July 6, 1956 she found a dead, decorticated
specimen, though with some color and fresh markings, of
Erronea caurica (LinNAEUS, 1758) (Plate 14, Figure
23) at Rancho Eureka, Punta Arena. The morphological
details of this specimen are: (42.6 23.0 18.5 18 16).
On July 7 Mrs. DuShane found a specimen of Staphy-
laea staphylaea (LinnaEus, 1758) (Plate 14, Figure
24) at Los Chilenos, a locality 4 miles beyond Rancho El
THE VELIGER
Vol. 12; No. 1
Tule, but before reaching Cabo San Lucas. The shell was
in excellent coridition, imbedded in a clump of fresh sea
weed deposited on the beach. The details of this shell are:
(20.0 12.0 10.0 24 20).
The largest and smallest shells available to me for
this study are listed to provide an estimate of size variation
found in the species. The 5 figures given are, in the fol-
lowing order: length, width, height (all in millimeters),
number of teeth on outer lip, number of teeth on columel-
lar lip.
ACKNOWLEDGMENTS
There are many people to whom I wish to express my
sincere thanks, in addition to those mentioned elsewhere in
this paper, for their contributions of material, field infor-
mation, and literature references; they are: the late Con-
rad Limbaugh, Emery Chace, Mrs. Helen DuShane, Mr.
and Mrs. Ben Purdy, Mrs. Twila Bratcher, Glen Bickford,
John Fitch, Lawrence Thomas, Gale Sphon, Mr. and Mrs.
Michael O’Brien, Mrs. Emily Reid (for the maps), Dr.
Takeo Susuki (for special processing of the photographs)
and, finally, Jean Cate for helpful suggestions and con-
tinued encouragement of my work as well as for the
photography for the plate illustrations.
LOCALITY INDEX
Many of the localities listed here were obtained from
handwritten labels in various collections. Every effort has
been made to verify the spellings, but in some instances
certain place-names were not found on any map or atlas
available to me; it is regretted #f any discrepancies occur.
1. Acapulco, Guerrero, Mexico
2. Agua Chale (approx. 24 mi S of San Felipe), E Baja
California del Norte, Mexico
3. ibid. (subfossil, Indian kitchen midden)
4. Albemarle Island, Galapagos Islands, Ecuador
5. ibid. (Banks Bay) ;
6. ibid., (Tagus Cove) (HERTLEIN, 1939; Pleistocene)
7. Algodones (Bahia), Sonora, Mexico (Tesora Exped.)
March 1966 :
8. Almejas (Playa), W Baja California del Norte, Mex-
ico (adjacent to Magdalena Bay)
9. Angel de la Guarda Island, Gulf of California
10. Anacapa Island, off Santa Barbara, California
11. Audencia (Bahia de), Colima, Mexico; Churea Ex-
pedition January 1962
12. Bahia Honda, Panama
Vol. 12; No. 1
THE VELIGER
Page 105
13. Bahia Salahua, 3 mi. N of Manzanillo, Colima,
Mexico (Las Hondas Hotel area)
14. Bahia Tenacatita (60 mi. N of Manzanillo, Colima)
15. Balboa Bay, Orange County, California
16. Balboa, Canal Zone, Panama
17. see no. 5, above
18. Barra de Navidad (30 mi N of Manzanillo), Jalis-
co, Mexico
19. ibid., first Churea Expedition, 7-11 January 1962
20. ibid., second Churea Expedition, February 1963
21. Bat Island, Costa Rica
22. Bacochibampo Bay, 1 mi N of Guaymas, Sonora
23. Bomber Beach, Puertecitos, E Baja California del
Norte, Mexico
24. Buena Vista, Las Palmas Bay (23°35’N Lat.; 109°
40’ W Long.), East Baja California del Sur
25. Cabo San Lorenzo, Ecuador
25a. Cabo San Lucas, Baja California del Sur
26. Cabrillo Beach, San Pedro, California
27. Caleta de Campo, Guerrero, Mexico; just N of
Acapulco
28. Cambria, California (subtidal rocks)
29. Cambria Radar Station, California (14 mi. offshore,
67 - 70 feet of water)
30. Cape San Lucas, S tip of Baja California del Sur
31. Caracol (Playa) near Bahia San Carlos, Sonora,
Mexico; Tesora Expedition, 29 March 1966
32. Cardalitos, Peru
33. Carmen Island, Marques Bay (offshore from Lore-
to), Baja California del Sur
34. Carmen Island, Salinas Bay, Baja California del Sur
35. Catalina Island, California
36. Cerralvo Island, Gulf of La Paz, E Baja Califor-
nia del Sur
37. Changame Island, Panama Bay, Panama
38. Charles Island, Galapagos Islands
39. Chatham Island, Galapagos Islands
40. Chatham Bay, Cocos Island, Costa Rica
41. Cholla Bay, Puerto Pefiasco, Sonora, Mexico
42. Clarion Island (Revillagigedos Islands), Mexico
43. Cleofas Island, Tres Marias Islands, Mexico
44. ibid., (Yellow Bluff)
45. Coaloil Point, Goleta, California (20 feet of water)
46. Clipperton Island (11°00’N; 109°20’W); 1958
47. ibid., (HERTLEIN, 1937)
48. Cocos Island (5°32’N; 86°59’W), Costa Rica
49. ibid., (INGRAM, 1951)
50. ibid., Chatham Bay
51. Concepcion Bay, East Baja California del Sur
52. Corinto, Nicaragua
53. Corona del Mar, Orange County, California
54. Coronado Island, W Baja California del Norte
55. Coyote Lagoon (Bahia), Concepcion Bay, E Baja
California del Sur
56. Creston Island, Mazatlan, Sinaloa; 22 Dec. 1960
57. ibid., under rocks, 25 feet of water, January 1964
58. Dana Point, Orange County, California
59. Desemboque (Playa), Sonora, Mexico
60. Enchura Bay, Peru
61. El Coyote Bay, within Concepcion Bay, E Baja Cali-
fornia del Sur
62. El Tule (Rancho), approx. 9 mi E of Cabo San
Lucas, Baja California del Sur, 22 February 1963
63. Elwood Beach, Santa Barbara County, California
63a. Empalme (approx. 10 mi. S of Guaymas), Sonora
64. Ensenada, W Baja California del Norte
65. Espiritu Santo Island, E Baja California del Sur
66. Flamenco Island, Panama Bay, Panama (under rocks
at minus tide; December 1960)
67. Galapagos Islands (00°01’S; 90°30’ W) ; Ecuador
68. ibid., HerTLEIN, 1939 (Pleistocene)
69. Gaviota Point, Mazatlan, Sinaloa; second Churea
Expedition, 22 January 1963
70. Geronimo Island, off Punta Baja, W Baja California
del Norte
71. Goleta, Santa Barbara County, California
72. ibid., Coaloil Point, on rocks, minus tide
73. ibid., Cookout Beach, University of California at
Santa Barbara
74. ibid., reef off Devereau School, N Goleta
74a. Guadalupe Island, Mexico (29°00’N; 120°00’ W)
75. Guanacaste, Costa Rica
76. Guaymas, Sonora, Mexico
76a. Gulf of Fonseca, Costa Rica-Nicaragua
77. Gulf of Nicoya, Costa Rica
78. Gulf of Montijo, Panama
79. Hood Island, Galapagos Islands
80. Indefatigable Island, Galapagos Islands
81. ibid., Academy Bay
82. Isabel Island, Galapagos Islands
82a. Isla Angel de la Guarda, Gulf of California
83. Isla de Malpelo, Colombia
84. Isla San Benito (Bahia de Sebastian Vizcaino), W
Baja California del Norte
85. Isla San Luis, E Baja California del Norte
86. Isla Santa Cruz, Galapagos Islands
87. James Island, Galapagos Islands
88. ibid., Sulivan Bay
88a. ibid., HERTLEIN, 1939 (Pleistocene)
89. Kino Bay, Sonora (approx. 75 mi. N of Guaymas)
90. Kobbee Beach, Canal Zone, Pacific Panama
91. La Jolla, San Diego County, California
93. La Libertad, San Salvador
94. La Paz, SE Baja California del Sur
Page 106
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95. La Penita, Nayarit (between Compostella and Puer- 140. ibid., Palos Verdes Point
to Vallarta) 141, Panama, Pacific Central America
96. Las Animas (Bahia) mainland, W of Isla Angel de 142. ibid., Panama Bay
la Guarda, E Baja California del Norte 143. ibid., Panama Canal (cement wall within Pacific
97. Las Gaviotas Beach, Mazatlan, Sinaloa entrance)
98. Las Varas, Nayarit (out of Tepic, via Compostella ) 144. Pelican Point, Puerto Pefasco, Sonora
99. Lobitos, Peru 145. Perlas Island, Panama Bay, Pacific Panama
100. Loreto, E Baja California del Sur 146. Playa Almejas (adjacent Magdalena Bay), W Baja
101. ibid., (very small shells) . California
102. Los Angeles Bay (shore W of Isla Angel de la Guar- 147. see no. 31
da) E Baja California del Norte 148. see no. 59
103. Los Chilenos (8 mi. E of Cabo San Lucas), Baja Ca- 149. Playa Santa Maria (Cabo San Quintin), W Baja
lifornia del Sur California
104. Magdalena Bay, W Baja California del Sur 150. see no. 130
105. Manta, Ecuador 151. Point Conception, California
106. Manacora, Peru 152. Point Fermin, San Pedro, California
107. Manzanello (Boca Chamela) = Manzanillo, Colima 153. Point Loma, San Diego, Calif. (lower lighthouse)
108. Manzanillo, Colima 154. Point San Telmo (mainland landfall W of Isla Santa
109. ibid., Boca Chamela Cruz), E Baja California del Sur
110. ibid., 20 miles W of - 155. Point Vicente, Palos Verdes Peninsula, California
111. ibid., 100 miles S of - 156. Portuguese Bend, Palos Verdes Peninsula, California
112. Margarita Bay, La Paz, E Baja California del Sur 157. Puertecitos (approx. 180 mi S of California-Mexico
113. Maria Madre Island, Tres Marias Islands, Nayarit border), E Baja California del Norte
114. ibid., EmMerson & Op, 1963 (dredge, off Arroyo 158. ibid., 24 mi N of -
Honcho) 159. ibid., 28 mi S of -
115. ibid., Puerto Balleto 160. see no. 115
116. Maria Magdalena Island, Tres Marias Islands 161. Puerto Chileno (8 mi E of Cabo San Lucas), Baja
117. Mazatlan, Sinaloa, Mexico California del Sur
118. ibid., Yacht Club 162. Puerto Escondido (approx. 11 mi S of Loreto), E
119. ibid., 100 miles S of - Baja California del Sur
120. ibid., Venado Island 163. ibid., under rocks in the lagoon
121. Melique, Jalisco, Mexico 164. Puerto Grande, San Salvador Island, Galapagos Is-
122. Mendia, Sinaloa, Mexico lands
123. Miramar Beach, Guaymas, Sonora 164a. Puerto Libertad (approx. 60 mi N of Guaymas),
124, Mission Bay, San Diego County, California Sonora, Mexico
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126.
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128.
129.
130.
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132.
133.
134.
USI),
Modesto, Sinaloa
Monserrate Island, E Baja California del Sur (ap-
prox. 50 mi. S of Loreto)
Monterey Bay, California
Montijo, Gulf of Montijo, Pacific Panama
Morro Bay, California
ibid., Point Buchon
Narborough Island, Galapagos Islands
Negritos (Parinas), Peru
Newport Bay, Orange County, California
Newport Beach, Orange County, California
Norse Beach (Cholla Bay), Punto Penasco, Sonora
135a. Paita, Peru
136.
Sie
138.
139.
Paitilla Point, Panama
Palmito del Verde (approx. 50 mi. S of Mazatlan)
Palo Seco Beach, Canal Zone, Pacific Panama
Palos Verdes Peninsula, California
165.
166.
167.
168.
169.
170.
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174.
178.
176.
Puerto Madre, Chiapas, Mexico
Puerto Penasco, Sonora, Mexico
Puerto Vallarta (17 mi N of -), Jalisco, Mexico
ibid., small specimens from deep water
Pulmo Reef, (approx. 70 m S of La Paz), E Baja
California del Sur
Pulperia Reef, Panama Bay, Pacific Panama
Pulpito Point (approx. 40 mi S ofLoreto), E Baja
California del Sur
Punta Arena (approx. 70 mi N of Cabo Pulmo, E
Baja California del Sur
Punta Baja (approx. 15 mi S of El Rosario), E Baja
California del Norte
Punta Banda (approx. 15 mi S of Ensenada), W
Baja California del Norte
ibid., Cape (Todos Santos Bay)
Punta Canoas (approx. 55 mi S of El Rosario), W
Page 108
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Figure 2
Mauritia maculifera SCHILDER, 1932
Clipperton Island x1
. by
Figure 3 Figure 4
Mauritia depressa (Gray, 1824) Mauritia scurra retifer (MENKE, 1829)
Clipperton Island xX 14 Clipperton Island x 14
Figure 6
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SCHILDER & SCHILDER, 1939
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Clipperton Island DK:
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THE VELIGER
Page 109
176a. Punta Catedral, Costa Rica
177. Punto Colorado, Guaymas, Sonora
178. ibid., 1 mi S of -
179. Punta Diggs (6 mi S of Mazatlan), Sinaloa
180. Punta Dominical, Costa Rica
181. Punta Estrella (approx. 5 mi S of San Felipe), i Baja
California del Norte
182. Punta Final (S point of San Luis Gonzaga Bay), E
Baja California del Norte
183. Punta Parinas, Peru
184. Punta San Carlos (approx. 60 mi S of El Rosario),
W Baja California del Norte
185. Punta San Miguel (6 mi N of Ensenada), W Baja
California del Norte
185a. Punta San Roque (approx. 85 mi S of Cedros Is-
land), Baja California del Sur
186. Rancho Buena Vista (approx. 8 mi S of Santa Ro-
salia), E Baja California del Sur
187. Rancho El Tule (approx. 9 mi E of Cabo San Lucas),
Baja California del Sur
188. ibid., just E of -
189. Rancho La Cuevas (Popotle) (20 mi S of Tijuana),
W Baja California del Norte
190. Revillagigedo Island (off Mazatlan), Sinaloa
191. Salaverry, Peru
192. see no. 34
193. Salinas, Ecuador
194. Saladita Bay (near Guaymas), Sonora
195. Salsepuedes Island (near Tiburon Island), Gulf of
California
196. San Blas (Topolobampo), Nayarit, Mexico
196a. San Carlos Bay, near Guaymas, Sonora (approx.
8 mi N of -)
197. ibid., 7 mi NW of -
198. San Clemente Island, California (32°51’N; 118°30’
W)
199. San Felipe, E Baja California del Norte
200. ibid., 45 mi S of -
201. ibid., 50 mi S of -
202. San Francisco (Playa), near Guaymas, Sonora
203. San José Island (50 mi N of La Paz), E Baja Cali-
fornia del Sur
203a. ibid., Amortajarado Bay
204. San Jaunito Island, Tres Marias Islands, Nayarit
204. San Juanito Island, Tres Marias Islands, Nayarit
205. San Luis Gonzaga (approx. 85 mi S of San Fe-
lipe), E Baja California del Norte
206. ibid., on the reef
207. San Luis Island (approx. 70 mi S of San Felipe), E
Baja California del Norte
208. San Marcos Island (approx. 20 mi S of Santa Ro-
salia), E Baja California del Sur
209. San Miguel Island, California, 34°05’ N; 119°30’ W
210. San Lorenzo Island, just S of Isla Angel de la Guarda
211. San Nicolas Island, California, 33°15’ N; 119°30’W
Page 110
212.
DNB).
214.
Zoe
216.
2ilVe
218.
219.
220.
221.
222.
223.
224.
225.
226.
227.
228.
228a.
229}
230.
231.
232
233%
234.
230%
236.
2 3p
238.
239)
240.
Bes.
242.
243)
Deh
2p
246.
247,
248.
148s),
San Pedro Bay, Sonora, Mexico
San Pedro Breakwater, San Pedro, California
San Roque, Baja California
San Salvador Island, Galapagos Islands,
San Simeon, California
San Simeon, Chiapas, Mexico (20 - 30 fathoms, July
1961)
Santa Barbara Island, California (33°28’N; 119°02’
W)
Santa Catarina Landing (35 mi S of El Rosario),
W Baja California del Norte
Santa Cruz, Nayarit (10 mi. S of San Blas)
ibid., S end of Bahia Matanchen
ibid., Churea Expedition, 16 January 1962
Santa Cruz Island, California, 34°01’N; 119°50’ W
Santa Cruz Island, E Baja California del Sur, Mexi-
co (24°30’N; 110°45’ W)
Santa Cruz Island, Galapagos Islands, Ecuador
Santa Monica Breakwater, Santa Monica, California
Santa Rosalia, E Baja California del Sur
Santo Tomas Bay, W Baja California del Norte
Santa Ynez Bay (approx. 30 mi S of Santa Rosa-
lia), E Baja California del Sur
Sechura Bay, Peru
Seymour Island, Galapagos Islands
ibid., South - (HERTLEIN, 1959; Pleistocene)
Socorro Island, Revillagigedos Islands, Colima
Soldado Bay, N of Guaymas, Sonora; Paisano Expe-
dition, 30 April 1964
see no. 231 (except: 1939)
Taboga Island, Honda Bay (12 mi SW of Panama)
Tangola Tangola, Oaxaca, Mexico
Tagus Cove, Albemarle Island, Galapagos Islands
Tenacatita Bay, Jalisco; second Churea Expedition,
5-8 February, 1963
Tetas de Cabra (near Guaymas), Sonora
Three Arches Beach, Newport, California
Tiburon Island, Gulf of California
Topolobampo Bay, Sinaloa; second Churea Expe-
dition; 19 January 1963
Trailer Park, San Carlos Bay, Sonora; Tesora Expe- .
dition, March 1966
Tres Marias Islands, Nayarit; Tesora Expedition,
March 1966
Turtle Beach (12 mi S of Puertecitos), Okie’s Land-
ing), E Baja California del Norte
Valma Beach (2 mi S of Santa Barbara Pier), San-
ta Barbara, California
Venado Island, Panama Bay, Pacific Panama
Venado Island (off Mazatlan), Sinaloa; Churea Ex-
pedition, 21 December 1961
Vera Cruz, Panama
THE VELIGER
250.
ZF
2Oe.
253.
Zo
250%
256.
Vol. 12; No. 1
Vique Point, Panama Bay, Panama
Wenman Island, Galapagos Islands
White’s Point, San Pedro, California
Willard Bay (just N of Bahia San Luis Gonzaga), E
Baja California del Norte
ibid., Willard Point
see no. 118
Zoritos, Peru
SPECIES INDEX
ACQUINOCEALIS eerie 113, 114 SAD ELAMEXICONA cosviesssnsrssussisssnvee
AID UZINOSA rcssnssssnssrsiesinssesee 117 cepmcmensmnn JOB}, MG}, iG, ise
annettae ........... 104, 113, 115 MACUNLPETA cicerisisinissieinissienie 111
ATADICULA veresesssrmeineiseen 114, 115 MASSAUECNSIS carecsssstatsciresssrsein 114
ATENOSA vronseen 112 MONECtA) cece ee 117
AETICE PS crosses 103 nartaeformis .... von 117
barthelemyi 117 nigropunctata . . 114, 115
CapUtophidii recon 117 | Polynesiae
caputserpentis . 112, 117 POHTOG coerorsrrccoxteen income 118
PIG ETI an Oo 104 PUNCtUlALA .ecrrnreenienn 114, 116
Dr 11 TASALEIQNANA cracisressssnesesssseee 116
CORUUST ee 111 NE Baier Sr wi
CONETOVETSA cece 103, 113 rebar Lea a a ae
Gennes Mia Nine SCRUAETOTUT veiressisrsinsssnrseree 112
SGUTT Gene
AEPTESSA vesriseiren oe OB! TETIGD
SETNANAOENSIS eecescsrsniernenene 116 SOWENDYE cescrarn
FETTUZINOSA rcseirtisissinurrsinin SPAiC€D coc
ZEMMULA oeresorn staphylaea
hawaiiensis . tabescens .....
helvol pees. Leveson
IN CUTUGI eee transpiciens
DOC Nemecn ttn terrestres VENUSEA verneerensin
TSAD ELLA ereersessssssssssssssesees Uitell usm
1. Macrocypraea cervus cervinetta (KiENER, 1843)
Spéc. Icon. Coq., Cypraea, p. 72; plt. 2, fig. 1; plt.
Selig
(Plate 11, Figure 1)
Localities: 4 32 36 38 43 52 67 76 78 79 82 87 90
Largest shell:
Smallest shell:
94 99 104 105 106 112 113 116 117 120 122
135a 138 142 143 145 162 168 169 170 176a
1930 N97 2202225225023 023 a0 o Ome
254 255 256
97.1
ava fee
48.8
20.5
35.4
16.1
36 33
30 25
Shell variable in size, tending to be large; elongate ovate,
narrow for its length, strong, lightweight; humped ad-
apically, sloping in front, sub-cylindrical; aperture wide,
broadening sharply abapically; left margin rounded, right
Tue VELIGER, Vol. 12, No. 1 [C. N.CatE] Plate 12
Figure 7 Figure 8
Luria isabellamexicana (STEARNS, Luria isabellamexicana (STEARNS, 1893)
La Paz, Mexico x ft Clipperton Island xX 14
Figure 9 2 Figure 10
Zonaria a. annettae (DALL, 1909)
with Crepidula incurva (BRoDERIP, 1834)
Puertecitos, Mexico Xx 14
Zonaria a. annettae (DALL, 1909)
Puerto Escondido, Mexico x i
Figure 11 . : Figure 12
Zonaria a. annettae (DaALL, 1909), variant Zonaria annettae aequinoctialis SCHILDER, 1933
Puerto Escondido, Mexico K 2 Cabo San Lorenzo, Ecuador KE
Vol. 12; No. 1
THE VELIGER
Page 111
margin bluntly angled, neither more than noticeably cal-
loused; teeth long, strong, heavier on outer lip, extending
adaxially well onto columella from margin to base; base
convex, broadened outer lip flattened ; fossula short, crossed
with teeth, shallow or almost lacking; terminals heavy,
produced, openings wide, exaggeratedly so in front; pri-
mary shell color light grey, exposed through final covering
of light brownish-yellow as grey ocellae of various sizes;
a somewhat broadened light grey mantle line on right
dorsum; interior of shell light grey; terminals, base, lip,
and interstices bright brownish-grey; teeth dark brown;
semi-obscure small brow1 spots in base coloring.
This subspecies is associated with the Gulf of Mexico-
Caribbean Sea species Macrocypraea cervus cervus (Lin-
NAEUS, 1758). The Gulf of Panama seems to be the locali-
ty of greatest abundance for these animals. Macrocypraea
cervus cervinetta may have been separated from the orig-
inal stock and isolated by the Pliocene emergence of the
narrow land-bridge now connccting the Americas. Study
shows the two shell forms to be very close in growth, size,
color, and the living requirements of the animals them-
selves, Adult shells of subspecies range in size from quite
small to extremely large.
These animals appear to be algae feeders, living in
coral pockets and on ledges and on rocks and basalt sub-
strates. The species is fairly common, particularly in the
Gulf of Panama. Fossil records for the subspecies con-
sist, so far as I have been able to determine, of a single
specimen from the Pliocene at Seymour Island, Galapa-
gos Islands (DaLt & Ocusner, 1928).
2. Mauritia (Arabica) maculifera ScuitpER, 1932
Zool. Anz. 100 (7/8): 165
(Plate 11, Figure 2)
Localities: 46 47
Largest shell: B28 37.8 28.0 26 24
Smallest shell: 49.2 SHES 25.2 28 22
The shell description is omitted here, as the specimens
agree in color and morphological characters with the Ha-
waiian form (see CaTE, 1965) ; the shells are beach-worn
and a description would not correctly represent the spe-
cies. The shells I have seen from Clipperton Island seem
to be slightly smaller than the norm.
This animal has a broad living range that extends from
one side of the Pacific Ocean to the other without quite
attaining the American or Asian mainlands. With Clip-
perton Island the eastern anchor, it ranges to the Philip-
pine Island-Ryukyu Island-Japanese Islands axis in the
west; and north to south from the Bonin Islands possibly
to the Gilbert Islands just below the equator.
The 2 specimens listed above were collected by Conrad
Limbaugh in 1959. Other specimens of this species are
recorded in the literature (see HERTLEIN & ALLISON, 1960).
3. Mauritia (Arabica) depressa depressa (Gray, 1824)
Zool. Journ, 1: 77
(Plate 11, Figure 3)
Locality: 46
Shell data: 40.8 30.9 20.7 24 19
Shell fairly large for the species, short, wide, having a
flattened appearance, solid, heavy; terminals do not pro-
trude beyond periphery of marginal outline; margins
heavily, thickly calloused, angled and upswept, semi-
shouldered; base and outer lip of nearly equal breadth,
convex; aperture narrow, fairly straight, terminal open-
ings narrow; teeth of medium size and length, prominent;
primary shell-color off-white, overlaid with light chestnut-
brown, thickly ocellate, and divided the length of the right
dorsum with wide off-white mantle line; upper sides and
terminals light grey; margins and base off-white, with
sides, margins, and a narrow area of the base and outer lip
spotted with large, medium brown spots that become ob-
scure on the base; base, interstices, terminal interiors off
white, teeth brown.
The specimen listed here is a live-collected animal taken
at the low water line under a large rock just inside the
outer reef. Several dead shells were also collected among
the rocks on the beach at the same time (coll. Conrad
Limbaugh, 1959; pers. commun.). This species is fairly
well established in the islands extending across the north-
ern half of the Pacific Ocean, but has yet to be found in
the Hawaiian group. Reports of the species there have
proven to be Mauritia (Arabica) maculifera ScuipER,
1932. The addition of Clipperton Island extends the range
eastward from heretofore known localities.
4, Mauritia (Arabica) scurra retifer (MENKE, 1829)
Verz. Conch. Samml. Malsburg, p. 76
(Plate 11, Figure 4)
Localities: 46 47
Shell large, long, narrow, cylindrical, solidly formed;
terminals protruding, well developed; margins barely
thickened, rounded; base roundly convex; aperture long,
narrow, straight; teeth numerous, small, fine, not crossing
recessed columella; fossula broad, deep, a third the length
of columella, weakly dentate; primary shell color light
grey to grey-beige, dorsum overlaid with ocellate, light
chestnut-brown color which is bisected by a wide mantle
Page 112
line the length of the dorsum; terminals blotched with
black on either side; sides, margins, and most of base and
outer lip grey brown to smoky-brown, except for bright,
lighter coloring on center of base; sides, margins, and up-
per part of base marked with large dark brown spots;
remainder of base, outer lip, and interstices brownish-
beige; interior of shell, fossula, and columella off-white;
teeth dark brown.
Although small specimens are found occasionally, the
larger form seems to predominate. The live animals are
only fairly common; they are found living on rocks in a
very rocky area with active water flow. Dead shells on
the rocky beaches are in greater abundance. This species
appears not yet to have reached the American mainland
or its off-shore islands.
5. Lyncina vitellus polynesiae ScHILDER & SCHILDER, 1939
Proc. Malacol. Soc. London 23 (4): 187
(Plate 11; Figure 5)
Locality: 46
Shell data: 68.0. . 42.1 36.0 28 25
The single specimen listed here is a dead, subfossil shell
collected on the beach at Clipperton Island by Conrad
Limbaugh in 1959. Although it is in fair condition, it
does not reflect the true aspect or color of a live-collected
animal. However, the shell is large for the species, is typ-
ically formed, and, in this case, has the familiar golden,
glossy color of a subfossil shell; large white spots show
faintly through the dorsal golden color; sides, margins,
base, teeth, interstices, and interior of shell bright white;
teeth large, short and thick on outer lip; more numerous,
finer, not extending onto base, but longer adaxially, cros-
sing over columella and fossula; fossula broad, long, and
shallow; shell pyriform in outline.
This species is listed in the literature (HERTLEIN & AL-
LISON, 1960) as a single specimen found on a beach-flat
at the north side of the island. It would seem the species
is uncommon.
6. Lyncina schilderorum IREDALE, 1939
Austral. Zool. 9 (3): 303
Syn.: Cypraea arenosa Gray, 1824
(Plate 11, Figure 6)
Locality: 46
Shell data: 2 PAB NIGER AS Be
This specimen, very badly worn and decorticated from
beach rolling, was discovered in a bag of equally wom
Erosaria caputserpentis caputophidi ScuiLveEr, 1927, from
THE VELIGER
Vol. 12; No. 1
Clipperton Island. It was an exciting discovery and sub-
stantiates the report of this species at Clipperton Island
by HerTLEIN & ALLISON (1960). No shell description is
offered here because of the condition of the specimen; a
jagged hole penetrates the right dorsum abapically;.the
measurements and dentition given above are only approx-
imate. The species is undoubtedly rare at Clipperton Is-
land, but for that matter, it isnot at all common anywhere:
7. Luria isabellamexicana (STEARNS, 1893)
Proc. U.S. Nat. Mus. 16: 384; plt. 50, figs. 3, 4
(Plate 12, Figures 7, 8)
Localities: 4 14 18 27 30 36 42 43 46 48 50 76 79
94 95 98 108 109 110 111 113 117 160 169
172 190 204 232 244
Largest shell: 515 27.0 224 37 32
Smallest shell: 18.4 9.8 7.8 29 25
Shell cylindrically ovate, solid, strongly formed; dor-
sum flattened centrally, humped adapically, sloping sharp-
ly in front; base noticeably flattened adaxially, outer lip
likewise; aperture fairly straight, narrow; teeth numerous
on both lips, columella very fine, almost obscure, rudi-
mentary, short, lengthening somewhat onto central base,
not extending adaxially onto columella; lip teeth short,
weakly formed; fossula long, narrow, concavely shallow,
ribbed weakly with teeth (approximately 7 on adaxial
rim of fossula) ; terminals well developed, but only barely
protruding; margins thickened, barely angled; primary
shell color light grey, overlaid with light grey-brown in 3
wide, transverse bands, upon which are overlaid numerous
irregularly formed, broken narrow lateral black lines;
upper margins medium brownish-grey, fading into white;
base, teeth, interstices white; terminals bright, orange-
red; 4 dark brown spots superimposed on the orange-red
terminals in adult shells; sometimes the quadrimacula-
tion becomes a continuous brown band over the terminal
collars.
These animals are found living intertidally in coral for-
mations and on loose coral rubble at Clipperton Island; on
the West Mexican - South Baja California shores they oc-
cur on coral reefs; and at the Galapagos Islands in crevices
and under lava rocks. 5
A large population of this species has recently been dis-
covered at La Penita, Nayarit, Mexico; the shells, how-
ever, are very small on an average at this locality. Every
indication seems to point to Clipperton Island as being
the locality of greatest abundance of Luria isabellamext-
cana.
Most of the subspecies of Luria isabella isabella (Lin-
NAEUS, 1758) develop shells of a moderate size, seldom
THE VELIGER, Vol. 12, No. 1
Figure 13
Zonaria robertsi (HipaALco, 1906)
Kobbee Beach, Panama xX 2
Figure 15
Zonaria arabicula (LAMARCK, 1810)
Mazatlan, Mexico x 14
Figure 17
Bistolida r. rashleighana (MELvILL, 1888)
Clipperton Island X 2%
[C. N.Cate] Plate 13
Figure 14
Zonaria nigropunctata (Gray, 1828)
Galapagos Islands
Figure 16
Zonaria spadicea (SwaINson, 1823)
Point Vicente, California i
Figure 18
Bistolida t. teres (GMELIN, 1791)
Clipperton Island x 2
Vol. 12; No. 1
attaining the size of this eastern related species. The
morphological differences are outstanding, becoming in-
itially apparent in the eastern islands of Polynesia, espe-
cially at the Hawaiian Islands where it is recognized as
L. i, controversa (Gray, 1824).
As stated above, at La Penita a population of 203 (or,
possibly, more) animals of Luria isabellamexicana ‘were
discovered in the autumn of 1966 by Sergio Verboonen.
I examined 75 of these shells; all but a dozen were ex-
ceedingly small for the species; the largest shells range in
length from 30 to 36 mm, the remainder only from 18 to
22 mm.
8. Zonaria (Zonaria) annettae annettae (Dat, 1909)
Nautilus 22: 125
Syn.: Cypraea sowerbyi Kiener, 1845
Cypraea ferruginosa KiENER, 1843
(Plate 12, Figures 9 to 11)
Localities: 2 3 18 22 33 36 41 51 61 63a 65 76 82a
89 96 101 102 126 157 158 159 162 163 164a
166 169 172 181 194 195 196a 199 203 203a
205 206 207 208 210 228a 241 254
Largest shell: 44.0 24.0 19.4 26 19
Smallest shell: 21:4 18.5 10.9 19 14
Shell usually medium large, solidly formed, pyriform,
humped in back, sloping to the front; terminals produced,
semi-beaked, strong, well formed; margins thickened, no-
ticeably angled on left, weakly angled on right; aperture
fairly straight, wide, flaring somewhat abapically; teeth
short, well developed, finer on the columellar base; base
bulbously convex centrally, outer lip rounded; primary
shell color (observed in bulla stage) light grey, overlaid
by overlapping patterns of nearly solid chestnut-brown;
margins, terminals, base, and interstices medium chocolate
brown, with numerous irregularly sized dark brown spots;
teeth off-white.
This is a common species in the Gulf. The range for
this animal appears exclusively confined to the protected
waters of the Gulf of California; it seems not to occur
on the exposed southwest coast of Baja California. Its
habitat seems to be shallow water with algae covered rocks
and coral strata. The range of this species can be outlined
rather accurately by a line drawn across the Gulf from
Empalme (just S of Guaymas, Sonora) to a point roughly
midway between La Paz and Cape San Lucas (Baja Cali-
fornia del Sur), thence northward along the eastern shore
to San Felipe (Baja California del Norte) and across the
Gulf to Puerto Pefiasco; and finally south along the coast
back to Empalme. Zonaria annettae annettae is somewhat
THE VELIGER
Page 113
rare at San Felipe (Dr. Shasky, personal communication),
becoming more plentiful at Agua Chale and southward. I
have examined specimens from practically all the offshore
islands and can confirm them at the localities also (see
Cate, 1961).
Pairs of animals have been observed on several occa-
sions protecting clutches of small, amber-colored, gela-
tinous egg masses laid on rock or coral slab surfaces.
As noticed in other cowrie species, the larger of the two
animals present had its widely spread foot in contact
with the upper surface of the egg mass, while the smaller
remained immobile near by.
ScHILpER (1967) published a report of the Calyp-
traeidae found living attached to the shells of Zonaria
annettae annettae. This commensal relationship (see Plate
12, Figure 10) between the cowrie and a Crepidula is,
as far as I can ascertain, unique in the Cypraeidae. I
noted this phenomenon while collecting at Puerto Pefas-
co where I collected this species from under coral slabs
with at least one specimen of Crepidula incurva (BRovE-
RIP, 1834), and sometimes two, attached. A report (Mrs.
Helen DuShane, personal communication) tells of 9 spe-
cimens living on a single cowrie shell.
A subfossil specimen of Zonaria a. annettae I removed
from an Indian kitchen midden exhibits a 19.5 mm long
Crepidula scar in the dorsal surface.
It seems worth noting that the presence of Crepidula
incurva on Zonaria a. annettae seems restricted more or
less to the animals living within the triangle bounded by
Puertecitos, San Felipe, and Puerto Penasco; it does occur
also, but infrequently, on shells found at Guaymas and
Bacochibampo.
9. Zonaria (Zonaria) annettae aequinoctialis SCHILDER,
1933
Zool. Anz. 101: 193
(Plate 12, Figure 12)
Localities: 25 60 83 99 106 132 143 183 229 256
Largest shell: 46.2 29.3 234 18 13
Smallest shell: 40.9 26.4 21.5 17 12
Shell large, heavy, solid, widely ovate, subpyriform,
humped; terminals barely produced, strong, thickly
formed; margins thickly layered with nacre, rounded on
the left, subangled on the right; base bulbously convex,
outer lip rounded; aperture wide, curving left adapically ;
teeth short, interstices wide on outer lip and columella,
longer on the latter, crossing adaxially over columella
and fossula; fossula shallow, barely evident; basic shell
color light grey, dorsum overlaid almost solidly with ir-
Page 114
THE VELIGER
Vol. 12; No. 1
regularly applied chestnut-brown; marginal callus per-
pendicularly wide, sweeping up sides of shell, spotted with
large brown spots, some becoming obscure in the side
callus; callus, base, and interstices pinkish-lavender to
light-brown; teeth white. A small curious nacreous deposit,
free from spotting, is produced and forms the upper sur-
face of the terminal collar in front. This latter character
appears to be a constant feature in this subspecies, It was
present on 6 specimens I have examined.
This isolated subspecies of Zonaria annettae is rare as
we know it today, and appears to be restricted to the
coastal waters from Panama Bay (Canal Zone, Pacific
entrance) south to some as yet undetermined locality in
northern Peru. The specimens discussed here (Cate coll.
nos. C1467, C1468, C1469) are all dead beach shells,
one of which is in fairly good condition, the other 2 are
decorticated, revealing the 3 wide, brown transverse color
bands on the light grey primary color of the dorsum; all
shells are fully adult.
This subspecies is distinct and not difficult to disting-
uish from the nominate species living in the more northern
Gulf of California. The geographical separation between
the 2 forms appears complete. The larger, stronger shell
of Zonaria annettae aequinoctialis suggests the exposure to
the open sea these animals seem to prefer.
Because of the scarcity of animals and shells of this
subspecies, there is little information available on the dist-
ribution and habitat of it. I have seen only one live-col-
lected shell from a concrete abutment, covered with green
algae, at the Pacific entrance to the Panama Canal. It was
collected by Mrs. Ann P. Marti, Balboa, Canal Zone.
10. Zonaria roberts: (Hmatco, 1906)
Mem. Acad. Cienc. Madrid 25: 178
Syn.: Cypraea punctulata Gray, 1824
(Plate 13, Figure 13)
Localities: 52 75 76a 78 90 93 138 142 145 176a
5) esl}
Largest shell: 30.5 20.5 15.5 20 14
Smallest shell: I). 14.2 10.0 16 10
Shell of medium size, ovate, solidly formed, umbilicus
depressed; humped sub-centrally abapically; terminals
barely produced, well developed; margins thickly cal-
loused, barely upswept, subangled; base and outer lip
convex, base centrally swollen; aperture somewhat nar-
row, curving left adapically; teeth short, well defined,
crossing fossula but not columella; fossula triangular,
wider abapically, narrow, deep, inner margin prominently
dentate; primary dorsal shell color bluish-grey, with a
central transverse wide band of large brown blotches, all
overlaid with an irregular network of fine light chestnut-
brown lines and daubs of color; a median line runs the
length of the dorsum; sides light chocolate-brown, be-
coming lighter toward center of base and outer lip, densely
covered with small, semi-obscure dark brown spots; re-
mainder of base, lip, teeth, interstices, and interior of
terminals pale beige-white.
Aside from Panama Bay, probably less is known about
the range of this cowrie species than that of any of the
others in the Eastern Pacific group. This may be due to
the wild, inaccessible shoreline within its range, which is
recorded to extend from the central part of the Gulf of
California, where it is rare, south to the area of Paita,
Peru. It could exist within these boundaries, but I have
never seen specimens with verified collecting data from
north of La Libertad, San Salvador, nor from further
south than Panama Bay. The localities listed herein have
all been substantiated by responsible collectors. Ted Dran-
ga established the northermost locality listed in this paper
in 1953 at La Libertad; Mrs. Meralyn Mackley, Canal
Zone, Panama, has established the southernmost locality.
A report that the species occurs in the Galapagos Islands
remains unverified.
Mrs. Mackley sent me a box with 73 specimens she
had collected alive from under rocks at Kobbee Beach,
Canal Zone. Five of the specimens were Zonaria (Z.)
arabicula (Lamarck, 1810), while the other 68 were Z.
robertsi. It can therefore be considered to be common in
the general area of Panama Bay. There is a remarkable
similarity between the two species, Z. arabicula and Z.
robertsi, particularly in the dorsal aspect. An inspection of
the teeth, however, will assist in separating the two species.
The teeth of Z. arabicula are sharper, better defined; also
the aperture is longer, straighter, and narrower; more-
over, the shell is less pyriform than that of Z. roberts.
11. Zonaria (Zonaria) nigropunctata (Gray, 1828) —
Zool. Journ. 4: 81
Syn.: Cypraea trina KiENER, 1843
Cypraea gemmula WEINKAUFF, 1881
Cypraea massauensis SCHILDER, 1922
(Plate 13, Figure 14)
Localities: 4 5 6 38 67 79 81 88 88a 105 183 230
234 251
Largest shell: 39.3 22.0 17.2 24 16
Smallest shell: 20.5 11.3 8.7 21 13
Shell elongate, narrowly ovate, solid, strong, apex de-
pressed, humped adapically, sloping gradually to the
front; terminals clearly produced, distinctly formed,
thick, margins heavily calloused, shouldered and angled
THE VELIGER, Vol. 12, No. 1 [C. N.Cate] Plate 14
Figure 19 Figure 20
Monetaria moneta barthelemyi (BERNARDI, 1861) Erosaria helvola hawaiiensis (MELVILL, 1888)
Clipperton Island x 2 Clipperton Island XK Dy
Figure 21 Figure 22
Erosaria caputserpentis caputophidii SCHILDER, 1927 Erosaria albuginosa (GRAY, 1825)
Clipperton Island x 1g James Island, Galapagos Islands
Figure 23 19
Erronea c. caurica (LINNAEUS, 1758) Staphylaea s. staphylaca (LINNAEUS, 1758)
Punta Arena, Mexico x 1¢ Los Chilenos, Mexico xX 24
Vol. 12; No. 1
on the right side; aperture nearly straight, fairly wide;
teeth short, numerous, sharply formed, somewhat recessed
on columellar base, not crossing columella; fossula limited
to a narrow, shallow groove in first 3 teeth; base and lip
convex, more swollen centrally on base; primary shell
color medium grey, with 4 darker grey bands, all overlaid
with irregular pattern-network of pale yellow-brown; a
large dark brown blotch on either side of terminals, with
uncolored median surface separating them; sides, margins
pale beige, heavily overlaid with a massive concentration
of various sized spots and elongate blotches of dark
brown, becoming obscure toward center of base and outer
lip; terminal interiors, teeth, and interstices pale beige-
white.
Oxsson (1924) and Dati (1909) appear to provide
most of the presently available information concerning
the cowrie dispersal on the west coast of South America.
Otsson lists Zonaria nigropunctata from Lobitos, Peru;
DaLt, in a more general way, from Manta, Ecuador, to
Paita, Peru. The northern half of the mainland shore-
line is wild and difficult of access.
Mrs. Rose Burch (personal communication) states
that the mantle of Zonaria nigropunctata from the Gala-
pagos Islands is an almost tranparent pinkish beige-brown
with small darker spots (like fine sand), and projecting
from the mantle are delicate filaments, approximately
3mm in length; they are tipped with beige. The upper
surface of the foot is roughly the same color as the mantle.
Mrs. Burch states further that the animals have been
observed sitting on egg-masses at the end of the cold
season — October and November.
The species is only fairly common; its range is restricted
to the Galapagos Islands and a limited area on the Ecua-
dorian mainland to the east.
12. Zonaria (Zonaria) arabicula (Lamarck, 1810)
Ann. Mus. Hist. Nat. Paris 16: 100
(Plate 13, Figure 15)
Localities: 1 13 14 18 21 25a 36 37 44 51 52 69 76
77 79 80 90 99 106 108 114 115 116 117 137
142 167 169 176a 180 191 193 196 203 212
220 235) 236 238 247 255
Largest shell: 30.0 19.5 14.0 22 14
Smallest shell: 15.0 8.8 7.2 19 13
Shell of medium size, strong, varying from almost round
to narrowly ovate; margins thickened, flared semi-obtuse-
ly, concavely-angled above, edge irregularly indented,
large dark brown pits along edge of margin, terminals
barely produced, though well formed; base and outer lip
THE VELIGER
Page 115
convex, though appearing flattened because of lower plane
of aperture opening; aperture straight, narrow; teeth
short, longer on outer lip and abapical columellar side
of fossula; teeth sharp, knife-like, extending adaxially
across columella and fossula, strongly, thickly developed
cn inner margin of fossula, interstices deep; fossula long,
deep, narrow; primary shell color medium grey, thickly
overlaid with a close network of light chestnut-brown color
pattern; there is a narrow mantle line the length of the
central dorsum; medium greyish-beige sides are thickly
spotted with large dull brown spots; base, terminal inter-
iors, teeth, and interstices pale beige-white.
The species is fairly common, and, except for those
species ranging as far as Clipperton and the Galapagos
Islands, it has the broadest range of distribution of all
the Eastern Pacific cowries. Zonaria arabicula is collected
both intertidally and subtidally down to about 30 feet;
its range is from northern Peru to Guaymas, Sonora. It
is occasionally collected at locations on the southern half
of the east coast of Baja California south to Pulmo Reef.
There are unconfirmed reports of this species occurring
at the Galapagos Islands.
There is a parallel phenomenon in Zonaria arabicula
and the West Australian Zoila venusta (SowERBY, 1846) :
both species have evolved in midrange similar morpho-
logical variants. In both cases the shells develop to less
than 2 the normal size, even though they appear to have
attained maturity. The West Australian subspecies has
been named Z. venusta sorrentensis by SCHILDER in 1963.
The variant of Z. arabicula occurs in the area from Man-
zanillo, Colima, Mexico, to the Gulf of Nicoya, Costa
Rica. In both cases the species’ range is continuous, with
the variants and the “normal” animals all living together.
Fossil shells of Zonaria arabicula have been found in
the Upper Pleistocene (Grant & Gate, 1931, p. 753)
along the coast of Oaxaca, Mexico, a geographical center
that might be considered a point of origin for the species.
13. Zonaria (Neobernaya) spadicea (Swainson, 1823)
Tilloch’s Philos. Mag. 61: 376
(Plate 13, Figure 16; Plate 15, Figure 25)
Localities: 10 15 26 28 29 35 53 58 64 70 72 73 74
74a 84 91 124 127 129 133 139 140 149 150
Gi RD WS VSS MG} PS EE NG) altssee held
185a 189 198 209 211 213 214 216 218 219
223 226 228 246 252
Largest shell: 60.1 35.6 29.3 25 21
Smallest shell : 33.9 19.0 15}5) 22 18
Page 116
Shell large, subpyriform, solid, usually heavily formed;
dorsum evenly humped, base convex, outer lip noticeably
rounded ; aperture fairly wide; teeth short, barely reaching
onto columella or outer lip; fossula faintly formed or ab-
sent; terminals thickly, strongly formed; margins of med-
ium thickness, right side angled, rounded on left; primary
dorsal shell color light grey, base milk-white; central
dorsal surface to mantle lines on either side overlaid with
medium chestnut-brown, becoming darker, more intense
at mantleline edge; terminals light grey, base, teeth, and
interstices milk white.
This species ranges along the west coast of North Amer-
ica from Monterey Bay, California in the north to just
below Cedros Island, off the west coast of Baja California,
Mexico, in the south, a distance of approximately 800
statute miles.
The species is fairly common and occurs from the mid-
tide zone (often left exposed on and under rock ledges at
low tide) down to about 30 feet. The animals have been
observed living on various substrates including rocks, sand
and in the open on silty mud flats in shallow water.
Mrs. Twila Bratcher, Hollywood, states that on a dive
in from 8 to 20 feet at Puerto Santo Tomas Bay (Pacific
side of Baja California del Norte) in September the water
was extremely cold; that Zonaria spadicea was present in
great numbers crawling about on a silty mud substrate
and on rocks (14 specimens were counted on one rock) ;
all had their mantle withdrawn; the dorsa of some speci-
mens were covered with white spots of unknown origin;
there were no bulla stage individuals visible; the animals
on the rocks seemed to favor any depression available in
the rocky surface; when the translucent mantles were vis-
ible, they revealed peach colored spots; the foot was dark
brown.
Zonaria (Neobernaya) spadicea fernandoensis (ARNOLD,
1907) (Proc. U.S. Nat. Mus. 32: 558; plt. 50, figs. 8, 8a)
is probably one of the ancestors of the Recent Z. s. spa-
dicea. This fossil species from the Fernando Formation
of the California Pliocene has been obtained at Elsmere
Canyon, Newhall; Holser, Piru, and at Santa Barbara
and Deadman’s Island. John Fitch, San Pedro, has col-
lected subfossil specimens from kitchen middens on San
Miguel Island. The holotype is on deposit at the U.S.
National Museum in Washington, where it bears the cata-
logue no. 164961; the type locality is given as Elsmere
Canyon, 24 mi SE of Newhall, Los Angeles County, Cali-
fornia, Lower Pliocene. I am indebted to Dr. Joseph
Rosewater of the Smithsonian Institution, Washington,
for the photographs of this fossil (Plate 25, Figure 26).
THE VELIGER
Vol. 12; No. 1
14. Bistolida (Blasicrura) rashleighana rashleighana
MELviLL, 1888)
Journ. of Conch. 5: 288
Syn.: Cypraea rashleighana var. transpiciens
; Taytor, 1916
Journ. Conchol. London 15 (4): 122 - 123
(Plate 13, Figure 17)
Localities: 46 48
Largest shell: 22.5 13.6 10.9 19 19
Smallest shell: 19.7 12.6 9.6 18 18
The single specimen listed from Cocos Island, Costa
Rica, was collected by the 1905-1906 Expedition of the
California Academy of Sciences to the Galapagos Islands;
it is on deposit at this Institution. IncramM (1951, p. 155)
also recognizes this species at Cocos Island.
The 2 specimens, the meristic data of which are given
here, were found in the beach drift at Clipperton Island
by Conrad Limbaugh, La Jolla. They were collected to-
gether with Bistolida (Blasicrura) teres teres (GMELIN,
1791), on the rocky beaches. This, therefore, establishes
B. (B.) r. rashleighana as another cowrie species occurring
in the Eastern Pacific. But the species must be considered
as rare in either locality - Cocos Island and Clipperton
Island.
15. Bistolida (Blasicrura) teres teres (GMELIN, 1791)
Syst. Nat., ed. 13, no. 3405
Syn.: Cypraea tabescens Dittwyn, 1817
Cypraea punctulata Hwatco, 1907
(Plate 13, Figure 18)
Localities: 12 46 47 164
Largest shell: 34.6 19.5 15.0 25 27
Smallest shell: 23.0 11.9 9.3 23 26
Adult shells vary from large to small; shell subcylind-
rically elongate, strong; terminals protrude, well devel-
oped; margins thickened, heavily so on right side, angled,
sub-shouldered and uneven above with callous welts; aper-
ture gently curving, fairly wide; teeth numerous, longer
on outer lip, finer on columella and fossula, not ex-
tending onto base; fossula short, broad, barely concave;
base and outer lip convex; primary shell color very pale
bluish-grey, three-banded transversely with zigzag light
brown lines over which an irregular light brown dorsal
blotch is centrally superimposed; margins, terminals, base,
teeth, and interstices off-white; margins marked with
large brown spots, numerous on left side at widely spaced
intervals along right marginal shoulder.
THE VELIGER, Vol. 12, No. 1 [C. N.Cate] Plate 15
Figure 25
Zonaria s. spadicea (SWAINSON, 1823)
San Diego, California x1
Figure 26
Zonaria spadicea fernandoensis (ARNOLD, 1907)
Elsmere Canyon, Lower Pliocene x1
Vol. 12; No. 1
The species is fairly common at Clipperton Island; it
is collected both as live and as dead shells on the rocky
shore area. EMERSON & Op (1965) record a beach spe-
cimen (leg. Mrs. Angermeyer) collected at Puerto Grande,
San Salvador Island, Galapagos Islands; a second shell
was reportedly found at about the same time by another
inhabitant of the island.
16. Monetaria (Monetaria) moneta barthelemyi
(BERNARDI, 1861)
Journ. de Conchyl. 9: 48; pit. 1, figs. 4, 5
(Plate 14, Figure 19)
Localities: 46 48 79
Largest shell: Zbioll 2S 13.4 14 12
Smallest shell: 25.1 17.1 12.2 12 10
Eight shells were studied for this report. They were col-
lected by the Scripps Institution of Oceanography Expe-
ditions to Clipperton Island, 1956 and 1958. All 8
specimens are from Station B 4237 and, with the excep-
tion of one specimen, are dead, beach-worn and decor-
ticated; they are bleached white from the normal lemon-
yellow of live-collected shells. Lacking better shells, no
description is attempted here. HERTLEIN & ALLISON (1960,
plt. 22, figs. 1, 2) show that the species is variable in its
morphology. The same variability can be observed in
nearly any large population of this species; shell color will
also vary in intensity and hue. There seems no accepted
explanation of this phenomenon, which is noticeable
enough to have led some early authors to nomenclatural
excesses. Completely adult shells vary exceedingly in
size within a single population. Although this is a common
species at Clipperton Island in the intertidal area in shal-
low water (2 to 3 inches), it has, as far as we know, not
been found on the mainland.
A live-collected shell, obtained later, was given to me
by Conrad Limbaugh who stated that this species was
common on some of the tide flats at Clipperton Island.
Figure 19, Plate 14 is this specimen.
17. Erosaria helvola hawaiiensis (MELvILL, 1888)
Mem. Proc. Manchester Soc., Ser. 4, 1 (5): 226
(Plate 14, Figure 20)
Locality: 46
Largest shell: 28.2 19.8 14.8 19 15
Smallest shell: 15.6 12.0 8.1 16 11
Shell fairly large, broad, subpyriform; terminals only
barely produced, wide, well formed; margins thickened,
upswept, angled, pitted above; base convexly swollen,
outer lip rounded; aperture fairly straight, of medium
THE VELIGER
Page 117
width; teeth large, strong, lengthening onto outer lip,
longer at either end of columellar base, shorter centrally ;
fossula without dentition, long, narrow, shallow; primary
dorsal shell color light grey, numerously, thickly speckled
with fine white spots, all overlaid with less numerous,
larger light yellow-brown spots; sides, margins, base
rosy-beige, teeth and interstices pale cinnamon-orange.
The species is quite common at Clipperton Island where
it is found under coral rocks. It is also commonly seen as
dead shells on the beaches. The shells seem to be predomi-
nantly of the large size, although fully adult small shells
are in evidence as well.
18. Erosaria caputserpentis caputophidii ScuitpER, 1927
Arch. Naturgesch. 91/A (10): 108
(Plate 14, Figure 21)
Locality: 46
Largest shell: 3220 24.6 17.0 18 15
Smallest shell: 19.0 12.7 9.3 18 13
Shell wide, short, ovate, strong, solid, humped almost
centrally, though flattened in appearance; terminals not
produced; margins thick, sharply angled, noticeably up-
swept on either side; base and outer lip of approximately
equal width, both convex; aperture straight, curving left
adapically, narrow; teeth numerous, short, well defined,
not crossing columella, but front 2 or 3 teeth cross fossula ;
fossula narrow, shallow, a continuation of columella; pri-
mary dorsal shell color off-white to very pale beige, over-
laid witha loose network pattern of variously sized ocellae ;
narrow, winding mantle line on many shells; sides, mar-
gins, and half of base and outer lip dark blackish-brown;
a large greyish-beige area is superimposed over both front
and rear terminal; rest of base, outer lip and teeth off
white, interstices light brown.
Together with Monetaria moneta barthelemyt, this is
perhaps the most abundant cowrie species living at Clip-
perton Island. The animals may be found living under
coral slabs and out in the open in shallow water on shore
and reef flats. I do not know if this species ranges any
further eastward toward the mainland.
19. Erosaria albuginosa (Gray, 1825)
Zool. Journ. 1: 510
Syn.: Erosaria albuginosa nariaeformis ScHIL-
DER, 1930
(Plate 14, Figure 22)
Localities: 4 14 18 30 33 36 43 46 48 51 65 76 79
87 94 108 113 115 116 117 154 162 166 169
171 186 190 196 203a 204 224 228a 244
Page 118
Largest shell: 28.8 19.5 14.9 21 19
Smallest shell: 15.6 9.0 7.0 18 17
Shell relatively small, pyriform, sometimes ovate, strong,
humped; terminals produced, semi-beaked adapically;
base convex; aperture narrow, widening anteriorly; teeth
distinct, short, fine, more so on the base lip, not crossing
columella; fossula barely apparent or not at all; margins
thickened, sharply angled on both sides and noticeably
shouldered, more so on the right side; primary shell color
off-white, overlaid with pale yellow-brown which is bro-
ken with minute lacunae-like openings exposing small
white ocellae of primary color, upon which are numerous
larger medium-brown, irregularly sized spots with white
centers; margins, terminals, most of the base and outer
lip pale lavender; rest of base, lip, teeth, and interstices
white.
This is one of the two purely Eastern Pacific species to
range as far westward as Clipperton Island, the other
being Luria isabellamexicana. Like this latter species, the
animals seems to thrive in this remote eastern locality.
An Indo-Pacific species very closely allied to this, Erosaria
(Erosaria) poraria poraria (LINNAEUS, 1758), whose east-
ern range limit seems to be the Hawaiian Islands, ex-
hibits on the dorsum a coloring and marking pattern that
is almost identical with that of E. albuginosa. Yet the
two species are undeniably distinct.
Although it is difficult at this time to suggest a typical
locality for Erosaria albuginosa, I have assigned it to the
faunal area of the Gulf of California because of its
widespread distribution there. I have examined large series
of specimens from several other areas; none seem to pos-
sess enough morphological differences to separate them.
Many of the Clipperton Island shells have thickly formed,
sharply angled margins, especially on the right side. These
mostly beach collected specimens as well as live-collected
shells (leg. Conrad Limbaugh) were all in an excellent
state of preservation, but most of the specimens from
elsewhere do not have these characteristics. The develop-
ment of the shell margins at the Galapagos Islands ex-
hibits greater thickening. None of the specimens, and
particularly those from the Galapagos Islands that I have
examined seems to justify a separate taxon; Evosaria albu-
ginosa nariaeformis ScHILDER, 1930 is therefore here
considered a synonym.
LITERATURE CITED
ANTON, HERMANN EDUARD
1839. Verzeichnif§ der Conchylien welche sich in der Samm-
lung von Hermann Eduard Anton befinden. Halle, pp.
i-xvi; 1-110
THE VELIGER
Vol. 12; No. 1
ARNOLD, RALPH
1907. New fossil mollusks of California.
Mus. 32: 558; plt. 50
Bakus, GERALD JosEPH ae
1968. Zonation in marine gastropods of Costa Rica and species
diversity. The Veliger 10 (3) : 207 - 211 (1 Jan. 1968),
BRATCHER, TWILA
1965. A search for Cypraea isabella mexicana.
Shell News 13 (9) N.S.: 67
Cate, Crawrorp NEILL
1961. | Remarks on a variation in Cypraea annettae Daun,
1909. The Veliger 4 (2): 112-114; plt. 24 (1 Oct. 1961)
1965. Hawaiian Cowries. The Veliger 8(2): 45-61;
pits. 4- 10; 4 maps (1 October 1965)
CHACE, EMERY PERKINS
1958. The marine molluscan fauna of Guadalupe Island,
Mexico. _ Trans. San Diego Soc. Nat. Hist. 12 (19) : 319-332;
fig, 1 (16 October 1958)
Dati, WiLu1AM HEALEY
1909. Report on a collection of shells from Peru, with a sum-
mary of the littoral marine Mollusca of the Peruvian zoological
province. Proc. U.S.N.M. 37 (1704): 147-294: plts. 20
to 28 (24 November 1909)
Day, WituiaM HEALEY & WASHINGTON HENRY OcHSNER
1928. Tertiary and Pleistocene Mollusca from the Galapagos
Proc. U.S. Nat.
Hawaiian
(July 1965)
Islands. Proc. Calif. Acad. Sci., ser. 4, 17 (4): 89 ~ 136;
plts. 2-7
Darwin, CHARLES
1897. A naturalist’s voyage. Journal of researches into the
natural history and geology of the countries visited during the
voyage of H. M.S. ‘Beagle’ round the world. pp. 1 - 500
[chapter 17; Galapagos Archipelago]
De Roy, J.
1967. Shelling in the Galapagos. Hawaiian Shell News
15 (10) N.S. 94: 6 (Oct. 1967) and 15 (11) N.S. 95: 4
(Nov. 1967)
Emerson, WiLuiaM KeirH & WiLLIAM ERwoop Oxp, Jr.
1964. Additional records from Cocos Island. The Nau-
tilus 77 (3) : 90-92 (January 1964)
1965. | New molluscan records for the Galapagos Islands.
The Nautilus 78 (4): 116-120 (April 1965)
Grant, ULysses S., IV. « Hoyr Ropney Gate
1931. Catalogue of the marine Pliocene and Pleistocene
Mollusca of California and adjacent regions. Mem. San
Diego Soc. Nat. Hist. 1: 1 - 1036; 15 text figs.; plts. 1 - 32
(3 November 1931)
Guticx, Howarp E.
1965. _ A traveler’s guidebook.
California; 168 pp.; maps; illust.
HertTLen, LEo GEorcE
1937. A note on some species of marine mollusks occurring in
both Polynesia and the western Americas. Proc. Amer.
Philos. Soc. 78 (2) : 303 - 312; 1 plt.; 1 map
(December 1937)
Hertiein, Leo Grorce « ARCHIBALD McCLurE STRONG
1939. Marine Pleistocene mollusks from the Galapagos Islands.
Proc. Calif. Acad. Sci., ser. 4, 23 (24): 367 - 380; plt. 32
(10 July 1939)
A. H. Clark Co., Glendale,
Vol. 12; No. 1
THE VELIGER
Page 119
HertLein, Leo Georce « Epwin C. ALLIson
1960. Species of the genus Cypraea from Clipperton Island.
The Veliger 2 (4): 94-95; plt. 22 (1 April 1960)
INGRAM, WILLIAM Marcus
1948. The cypraeid fauna of the Galapagos Islands. Proc
Calif. Acad. Sci. 4" ser. 26 (7): 135 - 145 (28 June 1948)
1951. The living Cypraeidae of the western hemisphere.
Bull. Amer. Paleont. 33 (136): 1-170; plts. 1-4; 3 tables
(24 March 1951)
Krener, Lours CHARLES
1845, Spécies général et iconographie des coquilles vivantes
genre Cypraca. 1: 1-165; plts. 1-57 Paris, J.-B.
Bailliére
Lamarck, JEAN-Baptistz PrERRE ANTOINE DE MONET DE
1799. Prodrome d’une nouvelle classification des coquilles.
Mém. Soc. Hist. Nat. Paris 1; 63 - 91
Oxsson, AxeLt ADOLF
1924. Notes on marine mollusks from Peru and Ecuador.
The Nautilus 37 (4): 120-130 (April 1924)
ScHILDER, Franz ALFRED
1965. The geographical distribution of cowries (Mollusca:
Gastropoda) The Veliger 7 (3): 171-183; 2 maps
(1 January 1965)
1966. The higher taxa of cowries and their allies. The
Meligert9i (i) 3135 (1 July 1966)
1967. Calyptraeidae attached to living cowries. Hawaiian
Shell News 15 (12) N.S. 96: p. 5 (December 1967)
Srronc, Arcimpatp McCiure « G Darias HANNA
1930. Marine mollusca of Guadalupe Island, Mexico.
Proc. Calif! Acad. Sci. 4" ser. 19(1): 1-6 (4 June 1930)
1930. Marine mollusca of the Revillagigedo Islands, Mexico.
Proc. Calif: Acad. Sci. 4 ser. 19 (2): 7-12 (4 June 1930)
1930. Marine mollusca of the Tres Marias Islands, Mexico.
Proc. Calif: Acad. Sci. 4" ser. 19 (3): 13-22 (4 June 1930)
Page 120
THE VELIGER
Vol. 12; No. 1
The Cowrie Species Living at Guam
BY
CRAWFORD N. CATE
12719 San Vicente Boulevard, Los Angeles, California 90049
(Plates 16 to 25; 1 Map)
INTRODUCTION
IncRAM (1938)PUBLISHED A LIST of 18 cowrie species
from the Micronesian Island of Guam. His findings were
based upon collections made there in the years of 1923,
1925, and 1937. Almost all of the specimens are said to
have been collected in Tumon Bay. ScHILDER (1938/39)
identifies 33 species from Guam. In recent years (1956
and 1959) two separately executed mimeographed cowrie
lists have appeared on the island itself. Otherwise, at
various intervals only minor reference has been made to
the existence of cypraeids there.
It is the purpose of this report to bring up to date the
information we now possess about the cowrie family at
Guam. Listed will be all presently known cowrie species
living in the coastal waters of Guam; accurate collecting
station data on the individual species will be given; a
revised taxonomic system and nomenclature (fide ScHIL-
DER, 1966, 1968) will be used, and the species will be
illustrated for comparison and identification.
During a visit to Guam in November, 1966 I was able
to visit shell-collecting friends and examine their collec-
tions. I was able to obtain from these collections a fairly
clear picture of population densities for each species; my
estimates were later confirmed in conversations with the
collectors. With the exception of a number of species
that are very common at this island and at most Pacific
localities, there are about a dozen species that should
be called fairly common, with the remainder of the spe-
cies to be regarded as uncommon to rare.
Guam is located in Oceania (Micronesia) approxi-
mately 1500 miles east of Manila and some 3000 miles
west of the Hawaiian Islands, roughly at 14°00’ N Lati-
tude and 114°30’ E Longitude. It is the southernmost and
largest of the 15 recognized islands of the Marianas
Group, the northernmost group of islands and atolls in
Micronesia.
The island of Guam consists mostly of undulating
hills and sharp outcroppings of rock; a wide variety of
cypraeids is found in Guam waters, and the long coastline
provides a variety of ecological conditions for these ani-
mals, a point that I will enlarge upon below.
ECOLOGY anp HABITAT
The beach and its intertidal zone and the reefs provide
good shell collecting. At many points around the island,
notably at Two Lovers’ Point, the cliffs are steep, dropping
off precipitously to the shoreline below or directly into
the sea itself. These areas provide a habitat for the species
that require the more turbulent, aerated water for their
survival. Otherwise, as with most islands in the tropics,
the coasts and the bay areas are lined with reefs and coral
shallows.
The reefs vary in the amount of exposed surface at
low tide, some barely clearing the very tops, others drain-
ing free and exposing much marine life. The lengths and
widths of the reefs vary; they consist of both dead and
living coral, with emerging lava-basalt in evidence nearly
everywhere. This habitat is often sharp and jagged, and
covered with many species of algae; some substrates are
age-worn, trapping patches of sand into large and small
drifts; some reef areas in protected environments have
loose, movable rocks, boulders, and coral slabs, all of
which are the habitats of cowries.
The beach localities vary from broad, uncluttered sand
expanses to sand with rocky outcroppings to surfpounded
rough rock and coral rubble shelves. In the many bays are
tremendous low-tide expanses of very shallowly covered
(2 to 12 inches of water) porous sand on which many
of the more common cowrie species are crawling. Evosaria
caputserpentis (LinNAEuS, 1758) and Monetaria moneta
(Linnagus, 1758) seemed to be underfoot everywhere
in Agana Bay.
During my visit I was able to make a circuit of the
island and see the various habitats, collecting at some of
them. I was surprised at the warmth of the water.
Even though the list of presently known species is quite
extensive, there yet may be more deep water species to
be discovered in the future. For example, recently Bistolida
Vol. 12; No. 1
goodalli fuscomaculata (Pease, 1865) and Palmadusta
lutea lutea (GMELIN, 1791) were found at Guam for the
first time; the latter species was collected, by good fortune,
during my visit to the island, affording me an opportunity
to examine it with the living animal still in the shell.
In instances where quantities of available material
permit, I list the measurements (in millimeters) of the
largest and smallest specimen to indicate the size range
and variability of the Guam species. The formula of 5
numerical combinations, given at the beginning of each
species listing, indicates length, width and height of the
shell with the other two numbers giving the number of
teeth on the outer lip and the number of teeth on the
inner margin of the columella-base. Under the heading
“Localities” a numerical reference is given to the accom-
panying “Locality Index.”
ACKNOWLEDGMENTS
It has been only in recent years that intensive attention
has been given to shell collecting in the Guam area.
Military and governmental employees constitute most of
the workers in this field. It follows that these people are
the source for most of the information contained in this
paper. I list in alphabetical order the workers most re-
sponsible for specimens and collecting information made
available to me for this report: Mrs. Phyllis Eliason,
Donald and Kay Hiatt, Rica and Floyd Miller, Louise and
Tom Montgomery, Lorraine and Kenneth Rhein, and
Herbert Ward, all of Agafia. I wish to express to them
all equally my deep appreciation for their eager and
generous help. My thanks go to Mrs. Emily Reid, Berke-
ley, California, for the fine map; and to Jean Cate for
constant help, encouragement, and suggestions; to Dr.
Takeo Susuki for assistance in processing film; and to
Dr. L. G. Eldredge, Department of Zoology, College of
Guam, for references to the literature.
LOCALITY INDEX
1. Anae Island 6. Agat Bay
2. Asan 7. Apra Harbor
3. Asan Point 10. Bile Bay
4. Adelup Point 13. Cabras Island
5. Agana Bay 14. Camel Rock (Asan)
THE VELIGER
15. Cetti Bay
16. Cocos Island
17. Facpi Point
20. Glass Breakwater
23. Hilaan Point Reef
24. Haputo Point
27. Inarajan Bay
28. Lajes Rock
30. Mangilao
. Merizo
35. NCS Beach
36. Neye Island
37. Nimitz Beach
40. Orote Point
Page 121
. Pago Bay
. Piti Bay
. Piti Bay Reef
. Ritidian
. Rizal Beach
. Tantapalo Point
(Orote Peninsula)
. Taogam Point
. Tarague Beach
. Tipalao
. Talofofo Bay
. Tumon Bay
. Umatac Bay
60.
USO Beach
64. Ylig Point (Ylig Bay)
65. Ypao Point (Ypao Beach)
SPECIES INDEX
AMELINGE eictescssntssrnssssirsinseinnn 128
annulus ... ael29
arabica... wn 1783
GN EU Sere LOD)
GSELLUS HO
aurantium .. . 124
aurora ..... un 124
beckitiane ee een es 130
bistrinotataye ee OO
caputserpentis
CATNEOA eseonsn
CAUTECONy Sener nanan
childrentqr en
CRINENSIS eccsssse
clandestina ....
Gribrariaupe
Cylindrica ne
dautzenbergi
Ae PTESSA vecsvnn
Cglantina mercer
CTOSAP een ea SO,
CNIONES ee 126
firmbriata ne: Peel YY)
fuscomaculata 128
Glob ulusmeeese ee 129
goodalli ...... 128
BU OCU tS ecccennennecnncr nse, 127
icluola ee eee 130
hinundods in eee lane 128
In dtCame ee ee ee el 124
LTT OTALG enn 5 ()
tab ell er tnne en 126
KUEN Tb ooevceosseee 128
labrolineata .. 30
leviathan nO
limacina ........ Li eee 131
listert
Luteane
MACUIPETA oerccssssinsin 123, 124
MEA PPA crrccroesn
margarita . :
MNGTIACR Se ees
MAUIANG omens 124
moneta .....
nucleus
onyx
QUUITI eae ne near
poraria ........
punctata
SCRUMETIANG virvecessicersenssnieee 124
SCAUACTOTUM recvscesssresseesssreiee 125
SECUNIA ra en ae 124
staphylacayee 130
SEO Gp ee 128
CLT ee terete te career rears eae 124
COTES ip erate PS ce tee 124
testudinanae ee 126
CETUS S aoc cent etna 124
Vent niCulus@na es 126
vitellus 02... 125
ZICZAC We Leen oes a 127
Page 122 : : THE VELIGER
RITIOIAN PT.
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HAPUTO f°",
Pipe 0
N.C.S.BcACH £.° |
HILAAN PT. &
REEF eo
AMANTES PT.
(TWO LOVERS PT.)
TUMON
YPAO PT... BAY Ff,
YPAO BEACH y
O
DEDEDO
CAMEL ROCK
GLASS
BREAKWATER
APRA
HARBOR
oroTe IPA CH IPEUG
PT.
TIPALAOT:.
7TAOGAM PT.
TANTAPALO PT:
NEYE 1° Ke.
RIZAL BEACH
AGAT
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O
SANTA RITA
/AGAT
NIMITZ
BEACH
ANAE 1.9
TALOFOFO
fe}
D
fo}
~ a
m
FACPI PT. & ;
TALOFOFO
One GUAM
CETTI BAYS
AJES ROCK ° ! @ Shale
LANES A UMATAC _—_— —— |
UMATAC BAY A MILES
BILE BAY). INARA JAN DD (A RAJAN
\ af BAY
Wolk 2; No! 1
THE VELIGER
Page 123
1. Mauritia (Leporicypraea) mappa (LinnagEvs, 1758)
(Plate 16, Figures 1, 3a, 3b)
Localities: 3 7 16 23 31 44 45
Normal shell measurements:
Largest shell: U2) 52.0 42.1 39 31
Smallest shell: 715.9 48.8 40.6 38 31
Apra Harbor dwarf variant:
Largest shell: 38) 31.6 26.8 37 37
Medium shell: 50.5 29.3 24.2 37 36
Smallest shell: 43.6 25.0 Plait 37 34
The species is uncommon, found only in a few restricted
localities. The Guam shells seem to have a color affinity
with the Philippine representatives of the species, posses-
sing more of the pinkish coloring and less of the plain
white to very pale beige base as seen in specimens from the
Ryukyu Islands and more southern Pacific localities. The
most outstanding feature is the size difference between the
specimens collected in Apra Harbor and those from else-
where on the island. There is a dwarf population, occur-
ring together with the larger, normal appearing forms,
in Apra Harbor, although the two forms apparently do
not occupy exactly the same habitat. Aside from the
small size there seems to be no other aberration; the shell
is normal in appearance except that it is narrower, the
teeth much smaller, finer, and more numerous on the
columella, giving the effect of elongation of the characters
normally attributed to the species. These shells are not just
of freak occurrence, for I have examined no less than 8
specimens; the 3 listed here are in my own collection
(Cate no. C3448). The species, though most commonly
encountered in Apra Harbor, is found on the underside
of coral and lava rocks, in small marine caves, and in
coral pockets in the reef areas at certain other Guam
localities.
2. Mauritia (Arabica) arabica arabica (Linnagvus, 1758)
(Plate 16, Figure 2)
Localities: 7 20 23 30 43 53 55
Largest shell: 51.4 33.0 25.0 28 27
Smallest shell: 37.8 24.3 Wee 2a 21
This is one of the very few reasonably abundant cowrie
species at Guam. It is more or less widely distributed in
the rocky reef habitats, being found most commonly high
up on the reef crowns. Apra Harbor seems to be the
local center of distribution, with a noticeable thinning out
of the populations elsewhere around the island.
3. Maunitia (Arabica) eglantina (Ductos, 1833)
(Plate 16, Figure 4)
Localities: 2 30 40 44 45 65
Largest shell: 50.0 29.6 24.0 35 33
Smallest shell: 41.0 23.4 19.0 34 30
This species is uncommon to rare. At present it is known
only from a few scattered localities and is sometimes mis-
taken for Mauritia a. arabica. The Guam shells are gen-
erally small for the species when compared with those
found elsewhere in the Pacific region.
4. Mauritia (Arabica) maculifera ScuitpER, 1932
(Plate 16, Figure 5)
Localities: 2 3 23 30 36 40 44 55 65
Largest shell: 58.8 37.0 28.1 28 27
Smallest shell: 3559 23.3 17.7 25 20
This is a species that seems to range in a more or less
restricted fashion throughout the breadth of the Pacific
region, with its eastern terminus in the Clipperton Is-
lands and Hawaii (see Cate, 1965, p. 57; plt. 8, figs. 26a,
26b) ; the northern half of the Philippines, Ryukyu Is-
lands, Japan, and the Bonin Islands support the species in
the west. These animals occur fairly commonly in the
coastal waters of Guam, perhaps most abundantly in
Tumon Bay. The shells do not attain as large a size as in
the Hawaiian form (not much is presently known about
the Clipperton Island shells, although they appear to be
smaller at that locality), but resemble them in most
other aspects.
5. Mauritia (Arabica) depressa (Gray, 1824)
Zool. Journ. 1: 77
(Plate 17, Figure 6)
Localities: 3 36 40 44 55
Largest shell: 41.1 29.6 21.0 22 19
Smallest shell: 41.0 29.4 20.8 24 19
This is an uncommon species; it approaches here the
limits of its northern Pacific range as we presently know it
in the central-west Pacific region. It is the form that is
often confused with Mauritia maculifera. The appear-
ance of the dorsal color pattern and design of M. depressa
is in many cases almost identical with that of M. maculi-
fera. Because of this it is necessary to consider the ventral
features of the shell for identification. From the base the
shell often seems to be more orbicular and more flattened,
Page 124
the coloring is uniformly almost white to a very pale beige
in some cases, with the teeth more even in length,
particularly the columellar teeth. In contrast, the teeth
of M. maculifera are irregular in length, becoming finer
and shorter on the front half of the base. This feature
and the dark blotch across the rear half of the base of
M. maculifera are probably the most important separating
characters in these very closely related species. I might
mention that the shells of M. depressa are almost never
as large as those of M. maculifera; the largest specimen in
my experience is approximately 44mm (from the Sey-
chelles Islands).
6. Mauritia (Arabica) scurra indica (GmEuNn, 1791)
(Plate 17, Figure 7)
Localities: 3 5 7 55
Largest shell: 40.1 19.9 16.5 44 34
Smallest shell: 31.8 1) at 42 30
This species is rare at Guam and is very seldom taken
alive. For many years it was known only from 2 shells
(according to H. Ward and E. Boyer, both of Agama) ;
one was beach-collected, the other live-taken. The latter
was collected in 40 feet of water in Agafia Bay. Since
these first shells were discovered, however, others have
been collected, though infrequently. Because these shells
have the more calloused base, a paler shell color and more
numerous spots on the sides and their shape generally
is more cylindrical and the size is smaller than these
characters are in the nominate species, it seems that the
Guam specimens belong to the subspecies under which
I have discussed them.
7. Mauritia (Mauritia) mauritiana (LinNAEUS, 1758)
(Plate 17, Figure 8)
Localities: 13 20 24 31 52 55 64 65
Largest shell: 91.6 66.6 45.0 25 23
Smallest shell: 90.0 66.4 44.7 24 23
These animals require highly oxygenated water; they
occur in an environment where the waves pound the shore.
Such habitats are encountered at Guam on the ocean side
of the Glass Breakwater, at Ypao Point, Haputo Point,
and in the Tarague area. In these localities the species is
quite common, occurring to a depth of 25 feet.
8. Talparia talpa (LinnakEvus, 1758)
(Plate 17, Figure 9)
Localities: 2 3 7 44 45 47 48 53 55 64
THE VELIGER
Vol. 12; No. 1
Largest shell: 64.1 32:5 27.0 42 36
Smallest shell: 59.0 ~ 324 269. — 47/38
These animals seem to have a life cycle that finds them
disappearing from the intertidal zone at times, to re-
emerge after an interval of time. Even so, the species is
not common. In this and some other Guamanian cowries
I am made aware of how much they seem to resemble the
Philippine and Ryukyuan species, both in size and ap-
pearance. There can be no doubt about the close similarity
of the shells from these localities; yet this same fact does
not apply in all cases. Some forms are missing here or
exist in noticeably different aspects.
9. Cypraea tigris schilderiana Cate, 1961
The Veliger 3 (4): 108; plt. 19
(Plate 17, Figure 10)
Localities: 2 3 7 10 16 17 31 35 43 53 54 55 60
Largest shell: 92.1 69.5 50.0 26 23
Smallest shell: 90.0 636 45.6 25 22
The Micronesian form of this subspecies, not quite
attaining the great size of the Hawaiian shells, neverthe-
less compares favorably with the latter. The general bulb-
ous outline of the shell, the concavity of the base, with the
length, shape, and arrangement of the teeth, and the
simple dark color-spotting upon a white background, are
all reminiscent of the Hawaiian shell (see Cate, 1965: p.
58; plt. 8, figs. 27a, 27b). It is a common species in Guam
waters and can be found in most reef areas from the low
tide line down to roughly 50 to 60 feet. Though they are
often scattered about as single individuals, they are also
frequently encountered in pairs or even in larger groups.
10. Lyncina aurora (Scuroter, 1789)
Syn.:Cypraea aurantium Gmeun, 1791
(see CaTE, 1966: p. 240; plt. 32, figs. la, 1b)
(Plate 17, Figure 11)
Localities: 1 7 17 28 40 50
Largest shell: 95.6 57.1 44.8 39 35
Smallest shell: 88.7 55.8 448 36 34
This is a rare species living in deep Guamanian waters.
These animals were unknown in this area, it seems, until
as recently as May or June of 1964, at which time An-
thony Elbo, Agama, collected the first specimen at Tanta-
palo Point in about 80 feet; subsequently 4 more were
found by Thomas Cruz, Agana, probably in the same
general locality; another live specimen was collected by
an unknown member of the United States Air Force. The
Vol. 12; No. 1
THE VELIGER
Page 125
most recent specimens were taken dead in deep water off
Lajes Rock by Mr. Herbert Ward, Agana.
The 5 live-collected shells from off Tantapalo Point,
Orote Peninsula, were living under huge basalt boulders.
The single live animal taken off Anae Island was found
in 45 feet in a small coral cave with a sandy bottom. In
color and outward appearance the shells would be difficult
to distinguish from those of the more southern parts of
the range of this species. The 2 dead shells collected off
Lajes rock were in about 70 feet of water.
11. Lyncina argus (Linnaeus, 1758)
(Plate 18, Figure 12)
Localities: 2 3 6 7 43 44 45 53
Shell data: 82.6 43.5 34.1 40 41
This is a rare species in Guam waters; I saw only an
occasional specimen in the collections there. One speci-
men was found in 80 feet in Pago Bay; another was ob-
tained from about 40 feet in a coral cave in the Asan-Piti
area; still another was from the Apra Harbor Reef; and
the specimen listed above (Cate coll. no. C1827) was
collected at Agat Bay. This species is rarely taken alive
here.
12. Lyncina lynx (Linnaeus, 1758)
(Plate 18, Figure 13)
iFocalitiess:2) 3) 0) 6) 7 15) 16 31935 44 45) 47 959 169
Largest shell: 61.0 34.2 Ds). 34 25
Smallest shell: 29.0 Apa 14.5 22 18
This is a common species here and usually can be en-
countered in most of the reef areas. The species is well
represented in most island collections. The large specimen
listed above was collected at Merizo, the small one at
Apra Harbor reef.
13. Lyncina vitellus (LINNAEUS, 1758)
(Plate 18, Figure 14)
Localities: 2 3 6 7 31 36 44 45 47 55
Largest shell: 62.5 39.0 34.8 31 24
Smallest shell: 31.6 19.5 16.4 23 19
This has to be listed as a common species in Guam, al-
though it is not at all abundant. It is represented in all the
island collections, but not in as large numbers as the
preceding species. It occurs in the reef areas in coral
indentations and algae-lined pockets.
14. Lyncina ventriculus (Lamarck, 1810)
(Plate 18, Figure 15)
Localities: 2 3 40 53 65
Largest shell: 59.6 39.5 30.0 27 22
Smallest shell: 47.5 33.0 23.6 24 21
This species is uncommon to rare and is presently
known from 4 feet of water at Shell Beach, Tipalao; it
has been collected at Asan also. Most of the shells at
Guam are of good size for the species, compared with
representatives from the Cook Islands and the Philippines.
The largest specimen listed above was collected at Orote
Point, the other in Apra Harbor (Cate coll. nos. C3473
and C3509).
15. Lyncina schilderorum (IREDALE, 1939)
(Plate 18, Figure 16)
Localities: 2 3 14 44 45 60
Shell data: 31.4 22.5 16.0 29 26
This species is rare in Guam waters, and is represented
mostly by dead beach specimens. The single shell listed
(Cate coll. no. C3494) was live-collected on the reef in
Piti Bay in 1965. Because of the scarcity of the species,
little is known about its general distribution at Guam.
16. Lyncina carneola carneola (LINNAEUS, 1758)
(Plate 18, Figure 17)
Localities: 2 3 5 7 14 35 36 44 45
Largest shell: 39.3 24.7 20.7 28 22
Smallest shell: 25.8 I) 13.4 25 20
Though only occasionally found, this species is relatively
well represented in the island collections that I saw; but
anyone to whom I talked referred to it as scarce. The shells
are typical of the species and are of medium size for it,
although there are many very small specimens with com-
pletely adult shells.
17. Lyncina carneola leviathan (ScHILDER & SCHILDER,
1937)
(Plate 19, Figure 18)
Localities: 2 3 4 7
Largest shell: 62.8 40.4 33.8 31 29
Smallest shell: 51.3 30.0 24.8 30 23
Page 126
This relatively rare species has only recently been rec-
ognized at Guam as being distinct from Lyncina carneola
carneola. This island locality may be near the western
end of its range of distribution, for the shells seem to
be not as large as those from the eastern part of the range.
The principal differences between the 2 subspecies appear
to be anatomical and in the size and weight of the shell
(Kay «& Weaver, 1963, p. 80).
18. Chelycypraea téstudinaria (LinNaEvs, 1758)
(Plate 19, Figure 19)
Localities: 2 3 5 31 47 55 65
Largest shell: 92.3 45.6 36.8 4S 43
Smallest shell: 89.4 45.1 35.2 43 41
The species is rare; the largest specimen listed above
was found in a submarine crater (World War II) off
Cabras Island, the smaller shell was live-collected in a
coral pocket off Asan Point. I saw the species represented
only in a few island collections.
19. Luria isabella (LinnaEvus, 1758)
(Plate 19, Figure 20)
Localities: 2 3 7 23 31 35 44 45 55
Largest shell: 29.0 16.3 13.6 36 27
Smallest shell: 18.4 9.5 78 36 23
The species is only fairly common. It lives in the algae
covered coral in deep crevices and is most active at night.
Shallow-water reefs are its main habitat.
20. Erronea (Adusta) onyx onyx (Linnagus, 1758)
(Plate 22, Figure 38)
Locality: 7
Shell data: 39.2 24.6 20.2 22 22
This species is found only on the south shore of Apra
Harbor and is very rare; it probably has reached here
the northeastern limit of its range of distribution. The
above specimen, collected prior to 1959 by B. J. Smith,
Agana, is in the Cate collection.
21. Erronea (Erronea) ovum (GmeEuN, 1791)
(Plate 22, Figure 39)
Localities: 7 44 45
Largest shell: 27.9 7h 13.8 iI) 15
Smallest shell: 27.1 17.0 13.3 13 18
THE VELIGER
This species heretofore has been mistaken for Erronea
errones (LINNAEUS, 1758) ; it was so labeled in some.of
the collections at Guam. The species is rare there and can
easily be separated from the just mentioned species by the
orange colored interstices on both lip and columella; the
shell also is more pyriform. These northern shells seem to
lack the dorsal blotch of brown seen on the shells from
New Guinea and the southern Philippine area.
22. Erronea (Erronea) errones (LINNAEUS, 1758)
(Plate 22, Figure 40)
Localities: 7 44 45
Shell data: 34.5 19.8 16.2 18 15
Known to be one of the more common species in the
western Pacific region, this species is rarely taken by col-
lectors at Guam; the shell listed here is from Apra Harbor
and was taken alive by an unidentified SCUBA diver;
it was found in very deep water at the mouth of the
harbor.
23. Erronea (Erronea) cylindrica (Born, 1778)
(Plate 22, Figure 41)
Localities: 7 23 44 45
Largest shell: 32.3 14.2 11.1 18 24
Smallest shell: 19.2 9.8 78 13 15
Although fairly common, the species is nevertheless
never abundant in Guam. The shells can be quite large
as shown above, but for the most part they more often
approximate the size of the smallest shell listed. It is also
found as a subfossil in Apra Harbor sand pits. |
24. Erronea (Erronea) caurica (Linnaeus, 1758)
(Plate 23, Figure 42)
Localities: 3 6 55 59 ae
Shell data: 34.8 17.8 14.5 16 16
The shell listed here is in the Cate collection and is
said to have been collected in 1955 by Mr. Herbert Watd,
Agafia. There is little doubt of its being rare in these
waters. Mrs. Phyllis Eliason and Mrs. Jean Allen, both
of Agafia, report the species from Guam;- SCHILDER’
(1938/1939) also lists the species from there, but I saw
no specimens in Guamanian collections in 1966.
Tue VELIcER, Vol. 12, No. 1
Figure 1
Mauritia mappa (LINNAEuS, 1758)
Asan
Mauritia mappa (LINNAEUS, 1758)
Apra Harbor xz
Figure 4
Mauritia eglantina (Ductos, 1833)
Piti Bay XX i
photographs by JEAN M. Cate
{C. N.Cate] Plate 16
Figure 2
Mauritia arabica (LINNAEUS, 1758)
Apra Harbor mS i
Figure 3 b
Mauritia mappa (LINNAEUS, 1758)
Apra Harbor x
Mauritia maculifera SCHILDER, 1932
Asan x 2
1 i i 3 5 aN és ie i
4 = t Sf : re y s) =
: ‘ ; ; \ i
2
/"s 1 y : '
:
: t mo
i J
» = ea ‘ ‘ : : H =
v . : = = t j z =
- : : pale rg) et tae . at
x = ie ‘ - ni = -
1 =~ 7 2 PA + me: a » Oe: | >
Tue VELIGER, Vol. 12, No. 1
[C. N.Care] Plate 17
Figure 7
Mauritia depressa (Gray, 1824) Mauritia scurra indica (GMELIN,
Orote Point x 4
Agana Bay OX igs
STELLA!
Ftc oN
Figure 8
Mauritia mauritiana (LINNAEUS, 1758)
Tumon Bay
Figure 10
Figure 11
Cypraca tigris schilderiana CATE, 1961
Lyncina aurora (SCHROTER, 1789)
Apra Harbor Anae Island Xs
photographs by Jean M. Cate
THE VELIGER, Vol. 12, No. 1 [C. N.CarE] Plate 18
Figure 12 Figure 13
Lyncina argus (LINNAEUS, 1758) Lyncina lynx (LINNAEUS, 1758)
Asan Point x4 Merizo
Figure 14 Figure 15
Lyncina vitellus (LINNAEUS, 1758) Lyncina ventriculus (LAMARCK, 1810)
Merizo oF Orote Point x 2
Figure 16 Figure 17
Lyncina schilderorum (IREDALE, 1939) Lyncina carneola carneola (LINNAEUS, 1758)
Piti Bay xX 12 Tumon Bay Xx 12
photographs by Jean M. Carte
Tue VELIGER, Vol. 12, No. 1 [C. N.CateE] Plate 19
Figure 18 Figure 19
Lyncina carneola leviathan SCHILDER & SCHILDER, 1927 Chelycypraea testudinaria (LINNAEUS, 1758)
Adelup Point DX Asan Point x 4
Figure 20 Figure 21
Luria isabella (LINNAEUS, 1758) Pustularia mariae SCHILDER, 1927
Hilaan Point Reef x Apra Harbor X 3
Figure 22 Figure 23
Pustularia bistrinotata SCHILDER & SCHILDER, 1937 Pustularia margarita (D1LtwyN, 1817)
Apra Harbor X g Piti Bay Sy
photographs by JEAN M. Cate
THE VELIGER, Vol. 12, No. 1 [C. N.Cate] Plate 20
Figure 24 Figure 25
Pustularia globulus (LINNAEUS, 1758) Pustularia childreni (Gray, 1825)
Apra Harbor xX 3 Apra Harbor X 3
Figure 26 Figure 27
Monetaria annulus (LINNAEUS, 1758) Monetaria moneta (LINNAEUS, 1758)
Agat Bay XX Oy Merizo a2
Figure 28 Figure 29
Naria irrorata (Gray, 1828) Erosaria becki (GAskoIn, 1836)
Asan x 6 Asan x 5a
photographs by Jean M. Carte
Tue VELIGER, Vol. 12, No. 1 [C. N.CateE] Plate 21
Figure 30 Figure 31
Erosaria labrolineata (GaskoINn, 1849) Erosaria helvola (LINNAEUS, 1758)
Piti Bay X83
Figure 32 Figure 33
Erosaria caputserpentis (LINNAEUS, 1758) Erosaria poraria (LINNAEUS, 1758)
N CS Beach x 14 Pago Bay xX 3
Figure 34 Figure 35
Erosaria erosa (LINNAEUS, 1758) Staphylaea staphylaea (LINNAEUS, 1758)
Asan DS See Apra Harbor x 25 }
photographs by Jean M. Cate
Tue VELIcER, Vol. 12, No. 1 [C. N.Cate] Plate 22
Figure 36 Figure 37
Staphylaea limacina (LAMARCK, 1810) Nucleolaria nucleus (LINNAEUS, 1758)
Asan xX 12 Apra Harbor XK @
Figure 38 Figure 39
Erronea onyx (LINNAEUS, 1758) Erronea ovum (GMELIN, 1791)
Apra Harbor Kes Apra Harbor xX 2
Figure 40 Figure 41
Erronea errones (LINNAEUS, 1758) Erronea cylindrica (Born, 1778)
Apra Harbor DX sity Hilaan Point Reef x1F
photographs by Jean M. Catz
THE VELIGER, Vol. 12, No. 1 {C. N.Carte] Plate 23
Figure 42 Figure 43
Erronea caurica (LINNAEUS, 1758) Erronea listeri (Gray, 1824)
Tumon Bay x 14 Apra Harbor xX 3
Figure 44 Figure 45
Notadusta punctata (LINNAEUS, 1758) Palmadusta asellus (LINNAEUS, 1758)
Asan x 4 Apra Harbor XS 3
Figure 46 Figure 47
Palmadusta clandestina (LINNAEUS, 1767) Palmadusta lutea (GMELIN, 1791!)
Apra Harbor XK B N CS Beach x5
photographs by Jean M. Catz
Tue VELIGER, Vol. 12, No. 1 [C. N.CatE] Plate 24
Figure 49
Palmadusta ziczac (LINNAEUS, 1758) Palmadusta gracilis (GaAskoIN, 1849)
Asan Point xX 2% Apra Harbor xX 3
Figure 50 Figure 51
Palmadusta fimbriata (GMELIN, 1791) Bistolida teres (GMELIN, 1791)
Piti Bay Reef x 4% Apra Harbor x 1%
Figure 52 Figure 53
Bistolida goodalli fuscomaculata (PEASE, 1865) Bistolida kieneri (Hiparco, 1906)
N CS Beach xX 4 N CS Beach xX 4
photographs by Jean M. Carte
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se
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Tue VeuicrEr, Vol. 12, No. 1 [C. N.Cate] Plate 25
Figure 54 Figure 55
Bistolida hirundo (LinNAEUS, 1758) Bistolida stolida (Linnagus, 1758)
Apra Harbor
Xx at Apra Harbor x 2
Figure 56 Figure 57
Cribrarula chinensis (GMELIN, 1791) Cribrarula cribraria (LINNAEUS, 1758)
Tumon Bay XX they Apra Harbor XxX 2f
photographs by Jean M. Cate
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Vol. 12; No. 1
THE VELIGER
Page 127
25. Erronea (Melicerona) listeri (Gray, 1824)
Zool. Journ. 1: 384
(Plate 23, Figure 43)
Localities: 7 44 45 60
Largest shell: 17.6 10.0 13 16 15
Smallest shell: 9.6 7.6 5.7 15 16
This species is uncommon to rare, with most of the
shells being collected in Apra Harbor; it is not found
very often anywhere.
26. Notadusta punctata (LinnaEvs, 1758)
(Plate 23, Figure 44)
eocalities: 2 3 7 23 44 45
Largest shell: 14.6 8.1 TS) 22 20
Smallest shell: 10.5 5.6 4.8 20 21
The shells of this species are small when compared with
those from the more western Pacific region populations.
The species is uncommon in Guamanian waters and is
rarely collected. The island collections appeared not to
have more than one or two specimens in them.
27. Palmadusta (Palmadusta) asellus (LinNAEUS, 1758)
(Plate 23, Figure 45)
Locality: 7
Shell data: 17.8 10.2 8.0 18 18
The shell listed above was collected circa 1955 by B. J.
Smith of Agama. It is now in the Cate collection. The
species must be considered very rare in Guamanian tidal
areas. As I do not have first-hand information concerning
this species, it will have to be considered as provisionally
from Guam.
28. Palmadusta (Palmadusta) clandestina (LINNAEUS,
1767)
(Plate 23, Figure 46)
Localities: 7 44 45
Largest shell: 2 9.6 7.8 22 17
Smallest shell: 15.1 8.1 6.7 22 17
Although almost every local collector has this species
in his collection, it is not a common species. It occurs on
the reefs and is oftcn found in the Apra Harbor - Asan -
Piti area.
29. Palmadusta (Palmadusta) lutea lutea (GMELIN 1791)
(Plate 23, Figure 47)
Locality: 35
Shell data: 11.4 6.5 os) 17 16
I had the pleasure of being in Agafia on the day this
first reported specimen was found on November 18, 1966
on the reef at NCS Beach by Floyd Miller. We were
visiting at his home upon the return from a collecting trip
and he handed me the shell, still containing the living
animal, for identification. Since then another specimen
has been collected in the same general locality by Mrs.
Phyllis Eliason. As can be seen from the data given here,
the shell is unusually small for the species, though it was
fully mature, with a normal color patter)
30. Palmadusta (Palmadusta) ziczac (LinnaEus, 1758)
(Plate 24, Figure 48)
Localities: 2 3 44 45
Largest shell: 18.8 11.5 9.0 17 16
Smallest shell: 16.1 10.0 8.1 16 15
This species is found only rarely, less than a dozen
specimens presently being known from this area. Most
shells are live-collected; however, the smallest one listed
above (Cate coll. no. C951) is a dead beach: Irift find
at Asan-Piti. According to Warp (1956) only 2 live-col-
lected specimens were in Guam collections at that time;
it is still a rare species.
31. Palmadusta (Purpuradusta) gracilis (Gasko1n, 1849)
(Plate 24, Figure 49)
Localities: 7 44 45
Largest shell: Wel OS) 8.2 16 15
Smallest shell: 13.4 8.0 6.6 16 16
Apia Harbor seems to be the local center of distribution
for this uncommon species. It is found on the reefs and
under coral rubble. I did not note any other island
localities for this species in the local Guam collections.
32. Palmadusta (Palmadusta) fimbriata (GMELIN, 1791)
(Plate 24, Figure 50)
Localities: 2 3 7 15 44 45
Largest shell: H2r2 7.0 ond, 18 18
Smallest shell: 11.8 6.3 4.9 18 17
Page 128
This species is not very common at Guam, but collec-
tions show it to be widely distributed among the island’s
reefs.
33. Bistolida (Blasicrura) teres teres (GMELIN, 1791)
(Plate 24, Figure 51)
Localities: 2 3 6 7 17 24 30 44 45 55
Largest shell: SRS 18.7 14.2 25 25
Smallest shell: 28.4 15.5 11.8 24 23
This species is fairly common at Guam with a wide
distribution among the offshore reefs.
34. Bistolida (Blasicrura) goodalli fuscomaculata (PEASE,
1865)
Proc. Zool. Soc. London 1865: 515
(Plate 24, Figure 52)
Localities: 35 44
Shell data: ied 6.7 5:3 21 17
Shell small, strong, ovate, humped; terminals well de-
veloped, barely protruding, distinctively formed; right
margin calloused, angled, semi-shouldered, with minute
indentations along marginal edge; aperture almost straight,
narrow; teeth numerous, very fine, short, barely reaching
onto outer lip, not at all on columella; fossula rudimen-
tary, barely evident, denticulate; primary shell color grey-
beige, finely speckled with chestnut-brown spots over dor-
sum, larger spots thickly concentrated along both margins,
with a large chestnut-brown dorsal blotch, and smaller
supporting color blotches; 4 large, well defined dark brown
spots adorn the sides of the terminal openings, two at
each end of shell (quadrimaculate); terminals, base,
teeth, and interstices grey-beige.
There are two synonyms for this species: Cypraea
adelinae Roperts, 1885 (Tryon, Man. Conch., 7: 168;
pit. 4, figs. 46, 47) and Cypraea dautzenbergi Hwatco,
1907 (Mem. Acad. Cienc. Madrid, Cypraea: p. 362).
The first Guam shell of this seldom seen subspecies was
collected by Mrs. Phyllis Eliason, Agana, at NCS Beach;
a second specimen has now been found at Piti Bay. This
subspecies is rare enough so that, with a fresh-collected
specimen at hand, it seemed desirable to redescribe the
shell. It is distinct from the nominate species, B. (B.) g.
goodalli (SowrrBy, 1832), more commonly encountered
at the Cook Islands in southwest Polynesia, which is
smaller, more slender, and does not have the brown ter-
minal maculations and the large, very dark dorsal blotch
of this subspecies.
THE VELIGER
Vol. 12; No: 1
35. Bistolida (Bistolida) kieneri (Hmauco, 1906)
(Plate 24, Figure 53)
Localities: 7 27 35 55
Largest shell: 134. 75° ° 69 ive amet
Smallest shell: 13:5 74 5.7 17 414
This species is apparently a relatively new addition to
the cowrie fauna of Guam, where it was unknown to
collectors prior to 1960. Since then, however, a number
of these animals have been collected, and dead shells have
been found in beach drift. The species remains uncommon
to rare.
36. Bistolida (Bistolida) hirundo (Linnaeus, 1758)
(Plate 25, Figure 54)
Localities: 2 3 7
Largest shell: AOR § OO WM ie
Smallest shell: 7 aye) 4.1. O18 14
This uncommon species occurs at the Apra Harbor reef
and at Asan reef. Its occurrence elsewhere at Guam is:as
yet unknown.
37. Bistolida (Bistolida) stolida (LinNAEus, 1758)
(Plate 25, Figure 55)
Localities: 2 3 7 23 35 44 45
Largest shell: 27.8 16.5 12.8 20 20
Smallest shell: 25.6 14.7 11-1 19 17
At one time this species was thought to be very rare in
Guamanian waters, but subsequent discovery and collec-
tions now place this form nearer the common group. Most
local collections contain from 2 to 3 specimens.
38. Ovatipsa chinensis chinensis (GMELIN, 1791)
(Plate 25, Figure 56)
Localities: 2 3 5 7 10 44 45 47 55
Largest shell: 340, 194 150) 1omuumls
Smallest shell: 31.0 176 138 18 18
This species, though uncommon, has a fairly wide distri-
bution on most of the island’s reefs. The morphological
characteristics of the Guam shells are. more closely re-
lated to the Philippine form, rather than to those of the
Hawaiian form (see Cate, 1965, 1966)
Vol. 12; No. 1
39. Cribrarula cribraria (LinNAEuS, 1758)
(Plate 25, Figure 57)
Localities: 2 3 4 7 44 45
Largest shell: 35.5 17.0 13.4 21 21
Smallest shell: 13.5 ici 6.1 il7/ 15
Apra Harbor and Tumon Bay are two well known
collecting localities for this uncommon species. It is not
found very often; it seems to prefer living deep within
the recesses, caves, and crevices of the reefs, and, being
more active nocturnally, it is rarely seen. See ScHILDER,
1967 (p. 103) for an explanation of the usage of the
generic name Cribrarula.
40. Pustularia (Annepona) mariae ScHILDER, 1927
(Plate 19, Figure 21)
Localities: 2 3 7 37 44 45
Largest shell: 17.0 il 10.3 43 27
Smallest shell: 13.1 8.8 8.2 32 21
This species is rare in collections in Guam and is repre-
sented entirely, I believe, by dead, beach shells only. The
localities listed here indicate a deep water habitat; practi-
cally nothing is known at present of the animal’s living
habits.
41. Pustularia (Pustularia) bistrinotata
SCHILDER & SCHILDER, 1937
(Plate 19, Figure 22)
Localities: 7 37 44 45 48
Largest shell: 19.2 758) 10.8 28 22
Smallest shell: 16.4 9.9 8.9 30 21
This species is uncommon; it is most often obtained as
a beach-rolled shell in which condition it has been fre-
quently mistaken for Pustularia margarita (Dittwyn,
1817), a species without dorsal pustules, a condition I
observed in the collections at Guam.
42. Pustularia (Pustularia) margarita (Di1LLwyn, 1817)
(Plate 19, Figure 23)
Localities: 7 44 45
Largest shell: 20.0 12.8 12.1 30 25
Smallest shell: 15.6 9.7 8.7 27 24
Like the preceding species, this one is uncommon in
Guam waters, with only an occasional shell being found
on the beaches, usually after a storm or high wind. The
specimens listed are from the Cate collection (larger shell),
and the Jean Kauanui collection (smaller shell). Both
THE VELIGER
Page 129
specimens are accompanied by labels giving Apra Harbor
as the collecting locality.
43. Pustularia (Pustularia) globulus (Linnazus, 1758)
(Plate 20, Figure 24)
Localities: 7 15
Largest shell: 17.8 11.2 10.0 36 26
Smallest shell: 17.3 10.5 9.3 33 29
This uncommon species is presently known only from
a few localities at Guam. While it is represented mostly by
dead, beach shells, occasionally a living animal is found.
Apra Harbor and Cetti Bay provide most specimens.
44. Pustularia (Ipsa) childreni (Gray, 1825)
(Plate 20, Figure 25)
Localities: 2 3 7 44 45 53 54
Largest shell: 22:3 15.3 12.5 41 25
Smallest shell: 14.9 10.0 8.0 33 22
A rare species at Guam, it is presently known only from
beach-collected specimens, obtained mostly after a storm.
As it apparently lives in deeper water, it is beyond the
reach of most local collectors.
45. Monetaria (Ornamentaria) annulus annulus
(Linnaeus, 1758)
(Plate 20, Figure 26)
Localities: 2 3 5 6 7 13 27 30 31 37 44 45 55 60 65
Largest shell: 21.1 16.2 11.0 13 10
Smallest shell: 15.0 10.4 Ue) 11 9
This species, unlike the following one, is common, but
not really abundant. It is found both in the shallow water
coral hills and on the sand flats that are barely covered
with water at low tide.
46. Monetaria (Monetaria) moneta (LinnaEus, 1758)
(Plate 20, Figure 27)
Localities: 2 35 7 13 27 30 31 35 36 37 44 45 55 60 65
Largest shell: 26.5 17.0 12.2 15 12
Smallest shell: 16.8 eS 7.8 12 11
This species is almost ubiquitous on the shallow sand
flats, on the reefs, around coral rubble, and on rocks. We
found it to be plentiful in the intertidal areas of Agafia
Bay and it seemed to live in areas where there were no
other mollusks present, except for a single bivalve species,
which remains unidentified.
Page 130
47. Naria irrorata (Gray, 1828)
(Plate 20, Figure 28)
Localities: 2 3 44 45
Largest shell: 10.9 6.3 4.8 16N
Smallest shell: il 5.0 3.7 16 14
This relatively rare species is found in the Asan-Piti
area inhabiting dead coral heads that are subjected to
a continual pounding surf. The Guamanian shells seem to
be small and narrow for the species, but possess the normal
color, spotting and other morphological characters of the
species.
48. Erosaria (Paulonaria) beck (Gasxotn, 1836)
(Plate 20, Figure 29)
Localities: 2 3 7 44 45
Largest shell: 10.5 5d 4.9 20 17
Smallest shell: 8.8 5.0 4.2 17 17
As of June, 1959, only one live-collected specimen was
known at Guam, but in recent years an occasional speci-
men has been found. This rare species is represented in
only a few local collections. Apra Harbor and Asan are
the localities where it has been taken most frequently.
49. Erosaria (Erosaria) labrolineata labrolineata
(Gasxoin, 1849)
(Plate 21, Figure 30)
Localities: 2 3 7 44 45 60
Largest shell: eal 10.0 7.9 18 17
Smallest shell: 14.4 8.5 6.7 16 16
A very rare species, it is known mostly from dead shells
picked up in the beach drift. Only 2 or 3 live-taken speci-
mens, from USO Beach and Piti Reef, are known at
present.
50. Erosaria (Erosaria) helvola (Linnaeus, 1758)
(Plate 21, Figure 31)
Localities: 2 3 7 16 31 35 36 44 45
Largest shell: 21.6 17.2 12.0 17 14
Smallest shell: 14.5 9.4 7.2 15 11
Though this is a fairly common species, it is never seen
in any quantity. It occurs in the reef areas at certain
locations and is collected mostly at night.
THE VELIGER
Vol. 12; No. 1
51. Erosaria (Erosaria) caputserpentis caputserpentis
(Linnaeus, 1758)
(Plate 21, Figure 32)
Localities: 2 3 5 7 10 13 15 16 20 23 27 30 31 35
36 37 43 44 45 47 48 52 53 55 59 60
Largest shell: 33ml 25.9 16.4 17 15
Smallest shell: 24.0 18.2 12.3 15 14
As can be seen by the number of localities listed, the
species is common and ranges over most of the island’s
reefs and rocky areas.
52. Erosaria (Erosaria) poraria (Linnagus, 1758)
(Plate 21, Figure 33)
Localities: 2 3 7 35 36 43 44 45 65
Largest shell: 18.5 14.4 10.3 18 16
Smallest shell: 13.4 9.1 6.8 20 13
This fairly widely distributed species is found on the
reefs and, on occasion, in the beach drift; however, it is
not common wherever it occurs at Guam.
53. Erosaria (Erosaria) erosa erosa (Linnagus, 1758)
(Plate 21, Figure 34)
Localities: 235 7 13 15 27 31 44 45 47 52 54 55 60 65
Largest shell: ends) BBE) Wes 21 15
Smallest shell: 21.4 13.0 9.6 18 11
This is the most commonly found cowrie species at
Guam, where it inhabits all the reef areas, living on the
algae-covered coral and under rocks and coral boulders,
and even on exposed tidal sand flats.
54. Staphylaea staphylaea (LinNAEus, 1758)
(Plate 21, Figure 35)
Localities: 7 44 45
Largest shell: 20.6 1225) 10.0 20 23
Smallest shell: 18.9 11.7 9.3 22 16
At present this uncommon species at Guam seems to
to be restricted in its distribution to the central-western
Apra-Harbor-Piti-Asan area; only on occasion is the shell
found.
Vol. 12; No. 1 THE VELIGER Page 131
55. Staphylaea limacina (Lamarck, 1810) Cate, Criwrorp Newt continued
1967. The cowries of the Ryukyu Islands. The Veliger
(Plate 22, Figure 36) 10 (1): 13-41; plt. 3; 2 maps (1 July 1967)
Localities: 7 44 45
Largest shell: 28.2 16.8 14.5 25 24
Smallest shell: 25.0 14.0 6.8 25 24
The species is rare at Guam; when found, it is in the
Apra Harbor-Asan-Piti area on the reefs, curiously in
the same habitat as the preceding species.
56. Nucleolaria nucleus nucleus (LINNAEUS, 1758)
(Plate 22, Figure 37)
Localities: 7 30 51
Largest shell: 25.4 15.1 12.3 27 21
Smallest shell: 18.0 11.6 10.0 23 20
This rare species is found now and then in the reef
area of Apra Harbor; it is represented by only a very
few specimens in the local island collections. The shells
seem to be generally smaller than those of Japan and the
Ryukyu Islands; the Philippine form also seems to be
shorter and broader than this shell.
LITERATURE CITED
Cate, CrawForp NEILL
1965. | Hawaiian cowries.
4-10; 4 maps
1966. Philippine cowries.
pits. 32-45; 3 text figs.;
The Veliger 8 (2): 45-61; plts.
(1 October 1965)
The Veliger 8 (4): 234 - 264;
1 map. (1 April 1966)
IncrAM, WILLIAM Marcus
1938. Cypraeidae from Guam The Nautilus 52 (1): 5-7
(July 1938)
Kay, EvizaBetH ALISON & Ciirton STOKES WEAVER
1962. Cypraeidae. Hawaiian marine mollusks 2 (20) : 75 - 78
(15 November 1962)
1963. Cypraeidae. Hawaiian marine mollusks 2 (21):
79 - 82 (1 February 1963)
1963. Cypraeidae. Hawaiian marine mollusks 2 (22):
83 - 86 (1 May 1963)
1963. Cypraeidae. Hawaiian marine mollusks 2 (23):
87 - 90 (1 August 1963)
1963. Cypraeidae. Hawaiian marine mollusks 2 (24):
91 - 94 (1 November 1963)
1964. Cypraeidae. Hawaiian marine mollusks 2 (25):
95 - 98 (1 February 1964)
ScHILDER, FRANz ALFRED
1965. Geographical distribution of cowries. The Veliger
7 (3): 171 - 183; 2 maps (1 January 1965)
1968. The generic classification of cowries. The Veliger
10 (3) : 264 - 273
ScHILpER, Franz ALFRED & Maria SCHILDER
1966. The size of ninety-five thousand cowries.
8 (4): 208 - 215
ScHILDER, Maria & FRANZ ALFRED SCHILDER
1967. Studies on East Australian cowries.
10 (2): 103-110; 9 tables
Eviason, PHYLLIS & JEAN ALLEN
1959. _A list of Guam shells. (privately circulated, unpublished
list of Guam mollusk species, including the cowries)
Warp, HERBERT
1956. (privately circulated list of Guam cowries; see remarks
for Palmadusta ztczac)
(1 January 1968)
The Veliger
(1 April 1966)
The Veliger
(1 October 1967)
Page 132
NOTES & NEWS
Notes on the Collection of
Tritonia festiva (STEARNS, 1873)
from the Seas of Japan
(Gastropoda : Nudibranchia)
BY
KIKUTARO BABA
Biological Institute, Osaka Kyoiku University
Tennoji, Osaka, Japan
(1 Text figure)
Tritonia festiva (STEARNS, 1873)
Latenbranchiaea festiva STEARNS, 1873. — Monterey Bay.
Tritonia festiva, Marcus, 1961. — Tomales Point.
Duvaucelia festiva, MacFaRLanp, 1966. — Monterey Bay.
Sphaerostoma undulata O’DonocuuE, 1924. — Vancouver Is-
land region; O’DonocHuE, 1926 (list).
Duvaucelia undulata var. muroranica Basa, 1940. — Muroran
(Hokkaido) ; Basa, 1957 (list).
Tritonia reticulata BercH, 1881. — Japan.
Here it is intended to give a summary of local informa-
tion gained from the study of our specimens of Tritonia
festiva which has, following MacFarLanpb (1966) priority
over T. reticulata Bercu, 1881. Possibly it has also priority
over Duvaucelia undulata var. muroranica. The study ma-
terial is listed below:
Specimens no. 1-4. Off Niigata, Japan Sea side of Japan,
70m (August 2, 1958, coll. by Dr. G. Kato).
Sp. no. 5. Sado, Japan Sea side, shore (May 9, 1956, coll. by
Dr. Y. Honma).
Sp. no. 6 - 10. Sado (May 20, 1966, coll. by Dr. I. Usuki).
Sp. no. 11. Abugashima, Toyama Bay, Japan Sea side, shore
(July 23, 1951, coll. by Mr. T: Abe and the author).
Sp. no. 12. Abugashima, shore (April 29, 1967, coll. by Mr.
T. Abe).
Sp. no. 13. Off Hayama, Sagami Bay, Pacific side of Japan
(Jan. 18, 1966, coll. by Biological Laboratory, Imperial
Household).
Sp. no. 14-16. Off Hayama (February 6, 1967, collector as
above).
These specimens ranged usually from 20 mm to 50 mm in
length, while the largest (specimen no. 14) showed the
maximum length of 75 mm in the preserved state. In
them the general ground colour of the back and sides
THE VELIGER
Vol. 12; No. 1
varied from a translucent whitish (specimen no. 11) or
pale yellow (specimens no. 6, 7) to as far as a deep
orange-red (specimens no. 5, 8 - 10, 13 - 16) or rather a
purplish red (specimens no. 1 -4 and 12). In every one
of the specimens there occurred prominent figures of
opaque white on the back. The maximum radular formu-
lae for the specimens dissected were 24 x 30:1-1:1-30
(specimen no. 11, body length 15mm), 35 x 40:1-1-1-40
(specimen no. 5, body length 20 mm), 45 x 80-1-1-1°80
(specimen no. 3, body length 35mm), and 50 x 90-1-1-
1-90 (specimen no. 16, body length 58 mm). The details
of the radular teeth and jaw-plates were as usual in the
genus Tritonia. The urn-shaped penis was proved to have
an apical papilla in the centre of the terminal disc (speci-
mens no. 3, 5, and 16). Nn
A conclusive statement may follow that our specimens,
though subject to variation in some respects, are possessed
of two of the eminent characteristics of Tritonia festiva
from the type locality: the first is the: presence’ of. the
opaque white figures on the back, and the second lies in
the formation of the apical papilla of the: penis. This
species, of which the type is known to be of a créam white,
has thus a distribution in the Pacific North America and
Japan, but so far there has been no record of collecting
of this form from the regions between the two.
ACKNOWLEDGMENTS
I wish to express my thanks to Dr. Genji Kato, Japan
Sea Regional Fisheries Research Laboratory; Dr. Yoshi-
haru Honma, Niigata University; Dr. Itaru Usuki, Sado
Marine Biological Station; Mr. Takeo Abe, Takaoka Se-
nior High School; and The Chief of the Biological La-
boratory, Imperial Household, for their generosity in
placing valuable collections made by them at my disposal
for comparative study. I wish to thank Mr. Clinton L.
Collier, San Diego, who kindly gave me facilities to refer
to some of the late O’DoNoGHUE’s papers on the nudi-
branchiate Mollusca from the Vancouver Island region. I
(on facing page —>)
Figure 1
Tritonia festiva (STEARNS, 1873)
A. Animal from above. Length 15 mm; specimen no. 11.
B. Left jaw from outside (40), specimen no. 11.
C. A half-row of radula (X200), specimen no. 11.
D. A half-row of radula (X50), specimen no. 16.
E. Distal part of male organ (X7), specimen no. 16.
a) penis; b) penis sac; c) apical papilla; d) vas deferens
F. Terminal disk of penis. Specimen no. 3.
a) opening of vas deferens
G. Terminal disk of penis. Specimen no. 5.
THE VELIGER Page 133
Vol. 12; No. 1
Page 134
am obliged to Mr. Steven J. Long, Pismo Beach, Califor-
nia, for a copy of the work by STEARNS.
LITERATURE CITED
Basa, KikuTar6 oo
1940. Some additions to the nudibranch fauna of the northern
part of Japan. Bull. Biogeogr. Soc. Japan 10 (6): 106 to
107.
1957. A revised list of the species of Opisthobranchia from
the northern part of Japan. Journ. Fac. Sci. Hokkaido
Univ. 6, Zool. 13 (1-4): 8- 14
Bercy, Lupwic SopHus RUDOLF
1881. Beitrage zur Kenntnif der japanischen Nudibranchien.
Il. Verh. k. k. zool.-bot. Gesell. Wien 31: 23 - 34
MacFarianp, Frank Mace
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
Marcus, ERNST
1961. | Opisthobranch mollusks from California. The
Veliger 3 (Supplement, pt. I) : 1-85; plts. 1-10. (Feb. 1, 1961)
O’DonocHveE, CHarves H.
1924. Notes on the nudibranchiate mollusca from the Vancouver
Island region, IV. Trans. Roy. Canad. Inst. 15 (1): 1 to
33; plts. 1, 2.
1926. A list of the nudibranchiate mollusca recorded from
the Pacific coast of North America, with notes on their
distribution. ‘Trans. Roy. Canad. Inst. 15 (2): 199-247.
STEARNS, RoBperT EpwArRDS CaRTER
1873. Descriptions of a new genus and two new species of
nudibranchiate mollusks from the coast of California. Proc.
Calif. Acad, Sci. 5: 77 - 78
Range Extension of
Tochuina tetraquetra (PALLAS, 1788)
to Hokkaido, North Japan
(Gastropoda : Nudibranchia)
BY
KIKUTARO BABA
Biological Institute, Osaka Kyoiku University
Tennoji, Osaka, Japan
Tochuina tetraquetra (Pauias, 1788)
Limax tetraquetra Patias, 1788. —Kurile Islands.
Tritonia tetraquetra, Bercu, 1879. — Unalaska (Aleutian Is.) ;
O’DonocuuE, 1922. — Victoria (Vancouver Island).
Tritoniopsilla tetraquetra, ODHNER, 1936.
Sphaerostoma tetraquetra, O’DonocHuE, 1926.
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Vol..12; No. 1
Duvaucelia tetraquetra MacFaRuanp, 1966. — Petersburg (Al-
aska) ; San Francisco Bay; Monterey Bay.
Tritonia gigantea Bercu, 1904. — Unalaska (Alaska).
Tritoniopsis aurantia Matrox, 1955. — Santa Catalina Island
(Southern California) ; Marcus, 1961.
In 1960 a single specimen of this notable species was col-
lected by Dr. Minoru Imajima, formerly a member of the
Shirikishinai Marine Biological Laboratory, Hokkaido
Kyoiku University, from 140 meters depth off Shiriki-
shinai near Hakodate, Hokkaido, Japan. As preserved, the
animal was greatly damaged, but it showed the charac-
teristics of the species in the thickly fringed rows of the
gills on the back-margins, and in having an extremely
large radula (90 X 220-1:220) consisting of a degraded
unicuspidate central and simply hamate (not filiform)
laterals. The total length of the body was more than
10 cm. From the above description it will readily be
seen that this species constitutes one of those forms which
cover in distribution the entire territory of the North
Pacific from east to west.
LITERATURE CITED
Bercy, Lupwic SopHus RUDOLF
1879. On the nudibranchiate gastropod mollusca of the north
Pacific ocean, with special reference to those of Alaska. Proc.
Acad. Nat. Sci. Philadelphia, pt. 1; 71-132; plts. 1-8.
1904. Nudibranchiata, Tectibranchiata-Pectibranchiata. In:
Semper, Reisen im Archipel der Philippinen. 9 (6), Lief. 1:
26 - 28
MacFar.anp, FRaNK Mace
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
Marcus, ERNST
1961. | Opisthobranch mollusks from California.
ger 3 (Supplement, pt. 1): 1-85; plts. 1 - 10.
Mattox, Norman T.
1955. Studies on the Opisthobranchiata: I. A new species of
the genus Tritoniopsis from southern California. Bull. S.
Calif. Acad. Sci. 54 (1): 8-13
OpuN_ER, Nits HjaLmMar
1936. | Nudibranchia Dendronotacea. A revision of the System.
Mélanges Paul Pelseneer. Mém. Mus. Roy. d’Hist. Nat.
de Belgique, Ser. II, Fasc. 3: 1057-1128; 1 plt.; text figs. 1-47.
1963. On the taxonomy of the family Tritoniidae (Mollusca:
Opisthobranchia). | The Veliger 6 (1): 48-52 (1 July ’63)
O’DonocHuE, CHARLES HENRY
1922. Notes on the nudibranchiate Mollusca from the Van-
couver Island region. III. Records of species and distribution.
Trans. Roy. Canad. Inst. 14 (1): 145 - 167; plts. 5, 6
1926. _A list of the nudibranchiate mollusca recorded from the
Pacific coast of North America, with notes on their distribution.
Trans. Roy. Canad. Inst. 15 (2): 199 - 247
Patras, P. S.
1788 Marina varia nova et rariora. Nova Acta Acad.
Sci. Imp. Petropolitana 2. St. Petersburg, 1784.
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Vol. 12; No. 1
THE VELIGER
Page 135
Current Paleontological Investigations
on Cenozoic Marine Mollusks
of the West Coast of North America
BY
WARREN O. ADDICOTT
U. S. Geological Survey, Menlo Park, California 94025
AND
SABURO KANNO
Tokyo University of Education, Tokyo, Japan
THIS REPORT 'SUMMARIZES recently completed studies and
work in progress on Cenozoic marine mollusks of the Pa-
cific coast of North America. Investigations are arranged
under several broad topical categories. Investigators are
listed alphabetically under each category together with
their institutional affiliation, if any, and current address.
Many studies span more than one of these but space lim-
itations have dictated against multiple listings. Certain
non-paleontological research such as isotopic studies re-
lated to age determination or paleoclimatic analysis is
included. Stratigraphic terminology is that of the individ-
ual workers. We are indebted to A. Myra Keen, N. FE Sohl,
and E. C. Allison for reading, and commenting on, the
manuscript.
TAXONOMY
Paleogene
Caroe S. Hickman (Washington State University, Pullman, Wash-
ington 99163) has completed a master’s thesis at the University of
Oregon on the molluscan fauna of the Oligocene Eugene Forma-
tion of western Oregon. Mrs, Hickman is currently preparing the
dissertation for publication.
Sasuro Kanno (Tokyo University of Education, Tokyo, Japan)
recently concluded an 11-month visit at the U. S. Geological Survey
in Menlo Park, California. He is completing a taxonomic-bio-
stratigraphic manuscript on the late Oligocene to middle Miocene
molluscan faunas of the Poul Creek and Yakataga Formations of
the Gulf of Alaska.
V. StanpisH MaAttory (University of Washington, Seattle, Wash-
ington 98105) is planning to have several of his recent students’
theses dealing with Paleogene mollusks of western Washington re-
written for publication in a museum-based monograph and novi-
tate series at the University.
1 Publication authorized by the Director, U.S. Geological Survey
A systematic study of Oligocene mollusks of the Pittsburg Bluff
Formation of northwestern Oregon is being carried out by ELLEN
J. Moore (San Diego Museum of Natural History, San Diego,
California 92112).
STanLey R. Primer recently completed a master’s thesis at the
University of California (Berkeley, California 94720) on the mol-
luscan fauna of the type Kirker Formation at Mount Diablo,
California. He has published a preliminary report on the biostrati-
graphy and correlation of the Kirker Formation.
Harotp E. Voxes (Tulane University, New Orleans, Louisiana
70118) has completed a manuscript on the molluscan fauna of the
upper Eocene and lower Oligocene Keasey Formation of north-
western Oregon.
Neogene
Warren O. Appicotr (U.S. Geological Survey, Menlo Park, Cali-
fornia 94025) has a paper in press on the taxonomy and biostrati-
graphy of early and middle Miocene gastropods from the south-
eastern portion of the San Joaquin basin, California. Included are
systematic descriptions and illustrations of more than 180 early and
middle Miocene gastropods.
Otuwareyisota S. ADEGOKE (Ife University, Ibadan, Nigeria) has
recently published reports on first occurrences of Swiftopecten,
Penitella, Platyodon, and Mya in the California Tertiary. His
doctoral thesis at the University of California is now in press; it
includes systematic descriptions of Neogene mollusks of the north-
western part of the San Joaquin basin, California.
Epwin C. Atiison (San Diego State College, San Diego, Califor-
nia 92115) continues his work on late Cenozoic mollusks of the
eastern Pacific, including a study of Cenozoic species of Arca.
Joun M. ArMENTROuT has completed a master’s thesis at the
University of Oregon (Eugene, Oregon 97401) ; this paper reviews
the molluscan fauna of the type Empire Formation (Pliocene) of
Coos Bay, southwestern Oregon, based on extensive collections in
the Department of Geology. Included in his study is an assemblage
of middle Miocene mollusks from previously unreported exposures
at Coos Bay.
EuceEnE V. Coan is preparing for publication his recently completed
doctoral thesis at Stanford University (Stanford, California 94305)
on late Cenozoic and modern Tellinacea of the northeastern Pacific
and arctic Alaska.
The long awaited manuscript on Pliocene mollusks of the San
Diego Formation, part of a series of reports on the Geology and
Paleontology of the San Diego, California area by Leo G. HeErt-
LEIN (California Academy of Sciences, San Francisco, California
94118) is ready to go to press. Utysses S. Grant, IV (University
of California, Los Angeles, California 90024) is co-author. HErt-
LEIN has begun work on a report on Pliocene mollusks of the Gala-
pagos Islands.
GrorcE L. KENNEDY (San Diego Museum of Natural History, San
Diego, California 92112) has completed a systematic study of
fossil Pholadidae of the eastern Pacific and is preparing a manu-
script for publication.
FE Stearns MacNem (5958 Prather Drive, Fort Myers, Florida
33901), formerly of the U.S. Geological Survey, is continuing his
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THE VELIGER
Vol. 12; No. 1
studies on Cenozoic mollusks of Alaska. He recently published a
monograph on the Cenozoic Pectinidae of Alaska.
Pliocene molluscan assemblages from the Imperial Formation near
San Gorgonio Pass, southern California, are being studied by
Micuaet A. Murpny and his students at the University of Cali-
fornia (Riverside, California 92507).
Ricuarp Rector, a student at the University of Washington
(Seattle, Washington 98105), has completed a thesis dealing with
mollusks of the Pliocene Quillayute Formation of the western
Olympic Peninsula, Washington, and is currently rewriting it for
publication.
Takeo Susuxki (University of California, Los Angeles, California
90024) continues his interest in middle Miocene mollusks of the
type Topanga Formation, southern California.
Pleistocene
S. Stirtman Berry (1145 West Highland, Redlands, California
92373) is continuing his descriptive studies of Pleistocene and
living warm water mollusks of the Gulf of California, western
Baja California, and southern California.
Wituiam K. Emerson (American Museum of Natural History,
New York, N. Y. 10024) is working on Pleistocene and. modern mol-
lusks of the Gulf of California and the Pacific coast of Baja Cali-
fornia. He has several recently published reports on gastropod
genera; many of these deal with the Muricidae.
GeorcE P. Kanaxorr (Los Angeles County Museum of Natural
History, Los Angeles, California 90007) continues his interest in
Pliocene and Pleistocene mollusks of southern California, concen-
trating, at the present time, on the molluscan fauna of the lower
Pleistocene Lomita Marl. His collections from the Pliocene San
Diego Formation have provided much of the material for Leo G.
HERTLEIN’s pelecypod monograph. J. ALDEN SUTHERLAND, also of
the Museum, has been assisting KaNnaxkorF in field excavation and
study of early Pleistocene mollusks from near Newport Bay, southern
California. SuTHERLAND has recently begun intensive collecting of
middle Miocene material from the Kern River area, central Cali-
fornia.
Jere H. Liprs (University of California, Davis, California 95616)
has several recent publications on the Pleistocene history and fora-
miniferal and molluscan paleoecology of insular faunas of the
Southern California Borderland; two of the reports were co-authored
by James W. VALENTINE of the same institution.
General
Ricwarp C. ALiison (University of Alaska, College, Alaska 99735)
is studying Cenozoic Turritellidae of the eastern Pacific. He and
Otuwareyisota S. ApEGOKE (Ife University, Ibadan, Nigeria)
have completed a report on a new early Tertiary turritellid genus
from the Pacific coast.
WituraM K. Emerson (American Museum of Natural History, New
York, N. Y. 10024) is continuing his studies on Cenozoic scapho-
pods of the Pacific coast.
A. Myra Keen (Stanford University, Stanford, California 94305)
has written up many of the pelecypod families for the Treatise on
Invertebrate Paleontology; the pelecypod volumes are now in press.
She has also summarized the Archeogastropoda and several super-
families in the Mesogastropoda and Neogastropoda for the second
gastropod volume of the Treatise. She has been assisted in this
work by EucEne V. Coan and other students.
Currorp M. Netson (Cabrillo College, Aptos, California 95003)
has begun work on a Ph. D, dissertation on Cenozoic Neptunea of
the eastern Pacific at the University of California, Berkeley.
Wits P. PopeNoE (University of California, Los Angeles, Califor-
nia 90024) is working on opisthobranch gastropods for the Treatise
on Invertebrate Paleontology. He is completing a joint study with
micropaleontologist RoperT M. KLEINPELL (University of Califor-
nia, Berkeley) on rates of evolution in tropical families in relation
to Lyellian correlation.
GerorcE E. Rapwin (San Diego Museum of Natural History, San
Diego, California 92112) is writing a monograph on the Columbel-
lidae of the eastern Pacific: ARNoLD Ross, of the same institution,
is engaged in a systematic study of ectocommensals of Cenozoic
mollusks.
Atiyn G. Smiru (California Academy of Sciences, San Francisco,
California 94118) is continuing his study of fossil chitons.
Jupiru S. Terry (Stanford University, Stanford, California 94305)
is preparing a manuscript on the cymatiid genera Argobuccinum,
Fusitriton, and Priene from her recently completed doctoral thesis.
She recently published a report describing the late cee genus
Mediargo.
WENDELL P. Wooprinc (U.S. National Museum, Washington, D.
C. 20560) has completed the manuscript for the last part of: his
four-part monograph on the geology and middle Eocene to early
Pliocene gastropods of the Canal Zone and adjoining parts of:
Panama (U.S. Geological Survey Professional Paper 306A-D).
BIOSTRATIGRAPHY
Warren O. Appicott (U.S. Geological Survey, Menlo Park, Cali-
fornia 94025) has a manuscript in press on Pliocene molluscan
faunas of the central Santa Cruz Mountains, California, and their
paleogeographic significance. He is working on molluscan faunas
from the upper Miocene Montesano Formation of southwestern
Washington, the type Temblor Formation of central California.
and lower and middle Tertiary formations of the Santa Monica
Mountains, southern California.
Otuwareyisoia S. ADEGOKE (Ife University, Ibadan, Nigeria) has
a manuscript in press, based on his Ph. D. thesis, on’ the molluscan
biostratigraphy of the classic Miocene and Pliocene sequences of
the Coalinga and Reef Ridge sections of the western portion of the
San Joaquin basin, California, originally studied by RALPH ARNOLD
in 1909.
KennetH W. Cirtacks and Aan R. Ormiston (Pan American
Research Laboratory, Tulsa, Oklahoma '74103) have been working
on the biostratigraphy of middle and upper Tertiary molluscan
faunas of the Gulf of Alaska and the Alaska Peninsula.
Eucene A. FrirscHE (San Fernando Valley State College, North-
ridge, California 91326) is completing work on a Ph. D. disserta-
tion at the University of California, Los Angeles, on the biostrati-
graphy and taxonomy of Miocene molluscan faunas of the Sierra
Madre Mountains, southern California.
Cuartes R. Givens (University of Georgia, Athens, Georgia 30601)
is adapting for publication his recently completed Ph. D. disserta-
Vol. 12; No. 1
tion at the University of California, Riverside, on Eocene molluscan
biostratigraphy of the Transverse Ranges, California. Givens has
published a preliminary report reaffirming the validity of the four
Eocene stages (Capay, Domengine, Transition, and Tejon) proposed
by B. L. Crank and H.E. Voxes in 1936.
Sapuro Kanno (Tokyo University of Education, Tokyo, Japan) is
describing the middle Tertiary faunal sequence of the Gulf of
Alaska based mainly on collections made during the summer of
1968 at Cape Yakataga.
Frank H. Kirmer (Humboldt State College, Arcata, California
95521) is continuing work on Neogene mollusks and associated
marine vertebrates of Cedros Island, Baja California.
Scotr McCoy, Jr. (Phillips Petroleum Corp., Bartlesville, Okla-
homa 74004) is studying the Tertiary molluscan biostratigraphy and
paleoecology of the Alaska Peninsula and Gulf of Alaska. He is
preparing a manuscript describing a new species of Pterynotus
from the middle Tertiary Poul Creek Formation.
Rosert G. McWitutaMs (Miami University, Oxford, Ohio 45056)
recently completed a doctoral dissertation on Eocene and Oligocene
biostratigraphy of central western Oregon including a thorough
re-evaluation of older work on marine mollusk faunas. He has
a manuscript in progress on mollusks of the Eocene Crescent For-
mation and is planning a report on mollusks of the type Twin
River Formation.
C. Percy Strone, a University of Washington graduate student,
has completed a manuscript on middle Tertiary mollusks and forami-
nifers from southwestern Washington.
Joun G. Vepper (U.S. Geological Survey, Washington, D.C.
20242) has recently published a detailed geological map of the
intertonguing marine and nonmarine Tertiary section of the
eastern part of the Caliente Range, California, and is now working
on the molluscan biostratigraphy in collaboration with continental
vertebrate and foraminiferal specialists. This study is expected to
provide greatly improved standards for provincial Miocene corre-
lation, VEDpDER is extending his field investigations to the nearby
portion of the Temblor Range and San Rafael Mountains. He is
also completing a biostratigraphic report on late Cenozoic mollusks
of the southwestern part of the Los Angeles basin, California.
Donato W. Weaver (University of California, Santa Barbara,
California 93106) and his students are preparing a summary report
on the geology and paleontology of the Channel Islands off south-
ern California. Included will be new biostratigraphic data on
Eocene and Miocene molluscan faunas.
PALEOECOLOGY
Joun W. Evans (Memorial University of Newfoundland, St.
John’s, Newfoundland, Canada) has been studying burrowing
mechanisms in pholadid pelecypods, in particular Penitella. He has
recent publications on the relationship of rock hardness to burrow
shape, identification of fossil burrows, and a method for estimating
rock hardness at time of burrowing.
Ciarence A. Hatt, Jr. (University of California, Los Angeles,
California 90024)) is studying the late Miocene and Pliocene
paleoecology of western San Luis Obispo County, California.
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Page 137
Joun P Kern (San Diego State College, San Diego, California
92115) has a manuscript in press on the early Pliocene molluscan
paleoecology of the eastern part of the Ventura basin, southern
California, based on a recently completed doctoral thesis at the
University of California, Los Angeles.
Rosert G. McWituiaMs (Miami University, Oxford, Ohio 45056)
is preparing a report on the paleoecology of late Eocene mollusks
and corals from near Seattle, Washington.
V. StanpisH Mattory (University of Washington, Seattle, Wash-
ington 98105), in cooperation with gcomorphologist Donatp J.
EasTERBROOK (Western Washington College, Bellingham, Washing-
ton 98225), is studying the paleoecology of late Pleistocene and
Holocene marine mollusks from glacio-marine tills in northwestern
Washington.
Louis MarincovicH expects to complete in mid-1969 a master’s
thesis at the University of Southern California (Los Angeles, Cali-
fornia 90007) on Pleistocene molluscan faunas of the higher
marine terraces at Palos Verdes Hills, southern California. To date,
42 previously unreported species have been found in the higher
terraces, many from new localities exposed during the continuing
urbanization of the area. W. H. Easton of the same institution is
collaborating with isotope geologist Joun K. Osmonp (Florida
State University, Tallahassee, Florida 32306) on a study of radio-
metric ages of molluscan faunas from these terraces. MARINCOVICH
is also working on the paleoecology of the early Pleistocene Lomita
Marl based on quantitative study of 119 species of mollusks.
Rosert FE Meape (California State College at Los Angeles, Los
Angeles, California 90032) has completed a doctoral thesis at the
University of California, Los Angeles, on the paleoecology of the
Santa Barbara zone of southern California. He has published pre-
liminary reports on the shallow water Pliocene mollusks that have
been incorporated in deep water turbidites of the Fernando Forma-
tion.
Gary RosENBERG has started a graduate research program at the
University of California (Los Angeles, California 90024) on the
paleoecology, distribution, and evolution of the bivalve mollusk
Chione in California.
Rosert W. Rowtanp has begun work on a doctoral dissertation at
the University of California (Davis, California 95616) on Pleisto-
cene mollusks of the Bering Sea. The study is being carried out in
cooperation with Pleistocene geomorphologist Davin M. Hopxins
(U.S. Geological Survey, Menlo Park, California 94025). Row-
LAND recently completed a manuscript from his master’s thesis on
the paleoecology of the molluscan fauna of the San Diego Forma-
tion of northwestern Baja California, Mexico.
Rosert J. STANTON, Jr. (Texas A&M University, College Station,
Texas 77843) has completed a paleoenvironmental study of Mio-
cene mollusks of the Temblor and Santa Margarita Formations from
the western side of the San Joaquin Basin near Coalinga, Califor-
nia. He has recent publications on the paleoecology of the upper
Miocene Castaic Formation of the eastern portion of the Ventura
basin, southern California.
Rosert R. TaLMancE (2850 Pine Street, Eureka, California 95501)
is studying the relationship of deep water mollusks from the Plio-
cene Rio Dell Formation of the Wildcat Group, northern Califor-
nia, to modern assemblages dredged off the nearby coast by trawlers.
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THE VELIGER
Vol. 12; No. 1
James W. VaLentTiNE (University of California, Davis, California
95616) is spending the 1968-1969 academic year on sabbatical
leave at Oxford University, England, where he is writing a book
on evolutionary paleoecology.
Joun E. Warme (Rice University, Houston, Texas 77001) com-
pleted a Ph. D. thesis at the University of California, Los Angeles,
on paleoecologic aspects of the modern ecology of Mugu Lagoon,
Ventura County, California. A recent publication concerning post-
mortem transport in fossil assemblages was developed from the
dissertation.
BIOGEOGRAPHY
Warren O. Appicott (U.S. Geological Survey, Menlo Park, Cali-
fornia 94025) has recently published reports on Oligocene and
Miocene zoogeographic discontinuities across the San Andreas fault,
California, supporting inferences of post-Oligocene cumulative
right-lateral slip of as much as 190 miles. He is currently completing
studies on Miocene latitudinal faunal gradients along the Pacific
coast and their relation to the San Andreas fault.
A recent publication by J. Wyatr DurHam (University of Cali-
fornia. Berkeley, California 94720) and FE Srearns MacNeEiIL
(5958 Prather Drive, Fort Myers, Florida 33901) summarizes mol-
luscan evidence for late Cenozoic marine migrations through
Bering Strait beginning in the late Miocene. More than 125 species
of Pacific origin have entered the Arctic-Atlantic area, whereas only
about 16 Pacific species seem to have had Atlantic origins.
A. Myra Keren (Stanford University, Stanford, California 94305)
is continuing her long range study of the relation of California
Miocene molluscan faunas to mollusks of the modern Panamic mol-
luscan province of Central America. She has begun preparations
for a revision of her important 1958 monograph on tropical mol-
lusks of the eastern Pacific in cooperation with JupirH S. Terry.
KATHERINE V. W. PaLMER (Paleontological Research Institution,
Ithaca, New York 14850) reviewed Tethyan affinities of marine
Eocene mollusks of North America, including the warm water
Venericardia planicosta and Velates perversus groups, in a recent
publication.
James W. VALENTINE (University of California, Davis, California
95616) has recent publications on the evolution of marine mollus-
can provinces and the climatic regulation of speciation. In coop-
eration with CLarENcE A. HALL, Jr. (University of California, Los
Angeles, California 90024), he is making a cluster analysis of late
Miocene molluscan assemblages from California as a means of
defining Tertiary molluscan provinces of the Pacific coast.
PALEOCLIMATOLOGY
Warren O. Appicotr (U.S. Geological Survey, Menlo Park, Cali-
fornia 94025) is studying middle and late Tertiary climatic change
in the nearshore marine environment through a distributional analy-
sis of warm water molluscan genera. He has completed a manu-
script on climatic change in the marginal eastern Pacific and is
working on a detailed study of Tertiary paleoclimates of the San
Joaquin basin, California.
Crarence A. Hatt, Jr. (University of California, Los Angeles,
California 90024) is studying growth layering in bivalved mollusks
as an aid in paleobiogeographic interpretation of eastern Pacific
molluscan faunas.
Rosert J. Stanton, Jr. (Texas A&M University, College Station,
Texas 77843) is working on a faunal and geochemical analysis of
Pliocene marine climate of the northwestern San Joaquin basin,
California, in cooperation with J. Rosert Dopp (Indiana Univer-
sity, Bloomington, Indiana 47401). A preliminary report is in press.
SYMPOSIA
Plans for a symposium volume on molluscan biogeography of
the eastern Pacific are being formulated by Victor A. ZULLO
(California Academy of Sciences, San Francisco, California 94118)
and Eucene V. Coan (Stanford University, Stanford, California
94305) . Several molluscan paleontologists will be invited to contrib-
ute to this effort.
Davin M. Hopkins’ (U.S. Geological Survey, Menlo Park, Cali-
fornia 94025) recently published symposium on the Bering Land
Bridge (1967) includes several papers containing Tertiary and
Quaternary molluscan faunal data from Alaska and Chukotka, in-
cluding a report by J. Wyatt DurHaAM and F Stearns MacNem
on Cenozoic migrations through Bering Strait.
Dwicut W. Taytor (Arizona State University, Tempe, Arizona
85281) is editing a review volume on malacology in western Amer-
ica. Contributions by molluscan paleontologists on marine ecology
and taxonomy are planned.
CATALOGS, GUIDEBOOKS
Warren O. Appicotr (U.S. Geological Survey, Menlo Park, Cali-
fornia 94025) is preparing annotated bibliographies of literature on
Neogene and Quaternary mollusks of the eastern Pacific Ocean
and arctic Alaska.
James H. McLean (Los Angeles County Museum of Natural
History, Los Angeles, California 90007) has completed a catalog
of living marine mollusks of Los Angeles County coast, California,
and is working on a distributional list of modern prosobranch gastro-
pods of the northeastern Pacific from Alaska to central Baja Cali-
fornia.
ELLEN J. Moore (San Diego Museum of Natural History, San
Diego, California 92112) recently completed a guidebook of fossil
mollusks of San Diego County, California, with illustrations of
Eocene, Pliocene, and Pleistocene species.
RADIOMETRIC AGE DETERMINATIONS
Several isotope geologists are working in cooperation with paleon-
tologists and biostratigraphers in attempting to provide radiometric
age control for Cenozoic stratigraphic sections along the Pacific
coast.
Vol. 12; No. 1
THE VELIGER
Page 139
Late Pleistocene and Holocene radiocarbon ages based upon
marine mollusks' shells have been reported by many workers during
the past 10 years or so. These have been particularly useful in
determining Holocene events in northwestern Washington and south-
western British Columbia, Canada.
In the past three years, uranium-series disequilibrium methods of
dating mollusk.shells have become increasingly useful in deciphering
late Pleistocene history of the Pacific coast during the past 200 000
years. H. HerBertT VEEH (Yale University, New Haven, Connecti-
cut 06511) and James W. VALENTINE (University of California,
Davis, California 95616), BARNEY J. Szaso and Joun N. RosHoLtT
(U.S. Geological Survey, Denver, Colorado 80225), Wittiam C.
BrapLey (University of Colorado, Boulder, Colorado 80302) and
Warren O. Appicotr (U.S. Geological Survey, Menlo Park, Cali-
fornia 94025), Horace G. Ricuarps (Academy of Natural Sciences,
Philadelphia, Pennsylvania 19103), and Dav L. TuurBer (La-
mont Geological Laboratory, Palisades, New York 10964), FE P.
FanaLe and O.A.ScHaArFFeR (Brookhaven National Laboratory,
Upton, New York 11973) have published papers during the past
three years dealing with the radiometric ages of Pleistocene terrace
faunas from California and southwestern Oregon.
Potassium-argon dating of Tertiary volcanic rocks and glaucon-
ites relevant to molluscan correlation is being carried out by JoHN
D. OxsrapovicH and by Donatp L. Turner, both of the U.S.
Geological Survey (Denver, Colorado 80225). OsrapovicH has been
dealing with Pliocene and Pleistocene ages as related to the Pacific
coast marine chronologies; TuRNER recently completed a doctoral
dissertation at the University of California, Berkeley, on K-Ar
dates concerning the Miocene foraminiferal chronology and has a
paper in press summarizing these data. He is planning to extend
his work, in collaboration with Warren O. AppicotT, to include
radiometric calibration of the Pacific coast molluscan sequence.
ISOTOPIC STUDIES
J. Rosert Dopp (Indiana University, Bloomington, Indiana 47401)
has completed a study of oxygen isotope and strontium paleotem-
perates of Pliocene and Pleistocene Mytilus from California. He
has undertaken a joint study with Ropert J. Stanton, Jr. (Texas
A&M University, College Station, Texas 77843) of oxygen isotopic
temperatures of Pliocene mollusks from the Kettleman Hill area
central California.
2
Cart L. Husss (Scripps Institution of Oceanography, La Jolla,
California 92037) is studying paleoclimatological aspects of a mixed
tropical and warm temperate late Pleistocene molluscan fauna from
Guadalupe Island, Mexico, by means of oxygen isotope determina-
tions. Epwarp C. Wirson (Los Angeles County Museum of Natural
History, Los Angeles, California 90007) has been studying the paleo-
ecology of this mixed fauna.
Heinz A. Lowenstam (California Institute of Technology, Pasa-
dena, California 91109) continues his oxygen isotope paleotem-
perature studies on Tertiary invertebrates.
OTHER STUDIES
J. Rosert Dopp (Indiana University, Bloomington, Indiana 47401)
has recently published a report on the relation of skeletal structure
and shell mineralogy to growth temperature and salinity based on
Pleistocene specimens of Mytilus from northwestern Baja California
and California.
GeorceE R. Crark II is completing a Ph. D. dissertation at Califor-
nia Institute of Technology (Pasadena, California 91109) on daily
growth increments in the Pectinidae. His work includes paleoeco-
logic applications in the study of late Tertiary pectinids from Baja
California.
H. Epwarp Cuirton (U.S. Geological Survey, Menlo Park, Cali-
fornia 94025), a sedimentologist, has been investigating the pre-
ferred orientation of disarticulated valves of late Cenozoic pelecy-
pods, particularly Psephidia, in the shallow water Elk River Forma-
tion of southwestern Oregon. In addition to field observation,
Cuirton has been introducing fossil shells into modern subtidal
environment and monitoring settling patterns by use of self-con-
tained underwater breathing apparatus. Similar studies, including
laboratory experimentation, have been made by Cortez W. Hos-
KINS (Union Oil Company Research Laboratory, Brea, California
92621) on a small, undetermined pelecypod that occurs abundantly
in deep water Pliocene turbidites in the Ventura basin, southern
California.
Page 140
THE VELIGER
Vol. 12; No. 1
THE CairorniA MALACOZOOLOGICAL Society, Inc.
announces
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Vol. 12; No. 1
THE VELIGER
Page 141
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Correction!
Due to an unfortunate error, the names of the shells
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were transposed. The species illustrated in Figure 3 is
Volva birostris (LinNAEuS, 1758) while that in Figure 5
is Volva brevirostris (SCHUMACHER, 1817).
Your Editor regrets the mixup and tenders his apologies.
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Page 142
METHODS & TECHNIQUES
A Simplified Vacuum Apparatus
for Collecting Small Nudibranchs
BY
J. SHERMAN BLEAKNEY
Department of Biology, Acadia University
Wolfville, Nova Scotia
(1 Text figure)
IF YoU HAVE EXPERIENCED the exasperation of collecting
small aquatic organisms with a combination of forceps,
eye dropper, screw-top jar, snap-lid vial, strainer and dip
net, then you will appreciate the simplicity of the appa-
ratus detailed in Figure 1. When a rubber sheet is attached
over the open end of the cylinder, finger pressure on the
rubber will displace air out of the tube at the side. If
this spout is placed in water in the proximity of a small
specimen and the pressure on the rubber released, the
specimen will disappear from its habitat and reappear
ay
™~
ik 3/32”
/ 5/16”
| 1/16”
Ome 54” Drill
Thickness - 1/16” : ;
Length - 134” Plexiglass Tubing
O.D.-34”
Thickness - 44”
Length - 114”
THE VELIGER
Glue Line
Plexiglass
EA ” x1 / 1 6”
~ Vol. 12; No. 1
instantaneously in the cylinder. The specimen can be
examined with a hand lens as it swims, crawls or attaches
itself within the transparent cage. Additional specimens
can be collected in the same cylinder as rapidly as your
finger can slowly depress and quickly release the elastic
cover of the container. This is a one-hand-only operation:
holding the cylinder and depressing the top with the index
finger. In cold water, neoprene diver’s gloves can be worn
without interfering with the efficiency of the technique.
Specimens can be transported to the laboratory in the
cylinder and stored in a refrigerator until examined under
a binocular microscope after removal of the rubber sheet.
Thus specimens quite literally need never be handled by
fingers or forceps during capture, transport, storage or
examination — not until they are removed for more de-
tailed examination or preservation. For this reason the
apparatus has been termed a Single Operation Collecting
Kit, abbreviated to SOCK, and respectfully designated an
“Acadian SOCK” in deference to an institution that has
fostered many a field biologist.
Some of the design presented in Figure 1 is rather
refined. The shut-off valve in the spout is not really
necessary because a vacuum effect limits the loss of
water. It is basically to keep specimens from escaping,
but they rarely find the opening, especially if the water
LD. 114”
O.D. 134”
|
|
3/16” |
Rounded Edges
1/16” Cut - 1%” Wide
~<— Plexiglass Tubing
O. D. 134”
Thickness - 14”
Length - 334”
~
=<— Plexiglass
1 ¥f," x 4”
Vol. 12; No. 1
THE VELIGER
Page 143
level is kept below the level of the spout. To lower the
water level in a SOCK, the diaphragm can be pumped
with the index finger and jets of water will be ejected
from the spout and no specimens, if they are crawling
mollusks. This apparatus was made specifically for col-
lecting sacoglossan and nudibranch mollusks in the size
ranges of < 2.5 cm. The O-ring and medium gauge den-
tal latex can easily be substituted by a string and a piece of
toy balloon or rubber glove. If you keep a few “Acadian
SOCKs” in your car, then a complete collecting kit for
sea shore or lake is always at hand and the expendable
parts can be purchased or replaced afresh in the shopping
center of even the smallest town.
My tide pool technique was to carry a basket loaded
with 10 Acadian SOCKs. Each species of nudibranch
collected was kept in a separate SOCK. If occasional
nudibranchs were too large or some of the dorids too
stiff then the rubber was flipped off and the specimen
dropped in with the others and the diaphragm replaced.
Specimens to be collected must be submerged, but it does
not matter if the SOCK has air or water or both in it.
The SOCK can be partly in air and water or totally sub-
merged. Specimens were scraped off the underside of
overturned rocks with the SOCK spout (or forceps) and
positioned in the aperture of the spout. The diaphragm
was depressed and the spout submerged in the nearest
pool of water, releasing the finger pressure at the same
time. The mucoid blob from the rock surface, now sub-
merged in sea water in the SOCK, would then assume
its natural shape and was examined with a hand lens.
The principle employed in the Acadian SOCK appara-
tus is applicable to larger and smaller diameter spouts
and cylinders. As long as the volume displaced by depres-
sing the rubber lid is greater than that of the spout, then
any small or delicate organism can be drawn back into the
cylinder within a gentle stream of water. I have field-tested
two sizes of SOCK apparatus in 7 countries since January
1968 and can attest to their effectiveness. In fact, I have
had the pleasure of using SOCK apparatus with SCUBA
gear in Zostera beds and in algal jungles. One need only
shake specimens off the plants or epizoans and then
leisurely “pick them out of the air” as they drift about;
or pieces of plants or hydroids or polyzoans with attached
nidosomes and adult nudibranchs can be broken off and
sucked up together.
The uninterrupted hours necessary to evolve the design
of the Acadian SOCK were made possible by a year of
Sabbatical leave from Acadia University, and a Canadian
National Research Council Senior Research Fellowship.
The author is further indebted to Professor George Hughes,
Chairman, Department of Zoology, Bristol University,
England, for providing workshop facilities where the pro-
totypes and present model were constructed.
Technique for Extraction and Mounting
of Gastropod Radulae
BY
GEORGE E. RADWIN
Curator of Marine Invertebrates
San Diego Natural History Museum
San Diego, California 92112
I HAVE USED the following procedure for extracting and
mounting radulae for a number of years. It is largely a
pragmatic technique developed by a process of trial and
error.
The extraction procedure differs depending on the size
and condition of the specimen from which the radula is
to be extracted. If a dried specimen is to be used, it
should be boiled in a saturated solution of trisodium phos-
phate (TSP). If the animal has dried deeply retracted
into the shell such boiling in TSP will usually cause sof-
tening and swelling sufficient to allow it to be seen and
removed from the shell. If the specimen is small, the
entire animal is usually treated. If a large animal is used,
the first step must be to remove the buccal mass, lying
immediately behind the foot and head areas and just
below the mantle cavity. The softened animal (or the
buccal mass) is then boiled in a concentrated solution of
sodium hydroxide for a short time(3- 10 seconds), re-
ducing it to a viscous brown film on the surface of the
NaOH solution. This brown scum is removed by pipette
and transferred to a watch glass with 70% ethanol. Gently
swirling the watch-glass, while carefully viewing it under
a dissection microscope will generally cause the brown
film to dissipate and leave the radula as a highly refractive
filamentous object. The radula is generally a ribbon-like
structure which may be removed by the aid of needles, and
stored in a vial of alcohol. If the radula of a toxoglossate
species [Conidae, Terebridae, Turridae] is extracted, the
radula may be a packet of tiny “darts.”
The radula under microscopic scrutiny is transferred to
a depression slide containing a drop of eosin stain and
allowed to remain there until the drop has dried. Using
a disposable hypodermic syringe the depression slide is
carefully filled with 70% ethanol. When the radula has
been sufficiently destained (5-10 minutes), the 70%
ethanol is removed, using the hypodermic syringe, and re-
placed with 95% ethanol. The lightly stained radula is
allowed to remain in the 95% ethanol for about 5 minutes,
then transferred with needles to a slide with a drop of
xylene or water-based mounting medium. Still viewing the
radula under the dissection microscope, a small section of
the radula is removed intact and segregated, the remainder
Page 144
THE VELIGER
of the radula being dissociated with a pair of very finely
ground needles. This step precludes confusion due to
overlapping teeth. If xylene is used, the radular material
is covered with a drop or more of canada balsam moun-
ting medium. If a water-based mounting medium is used,
the step involving xylene must be omitted and the radular
material is transferred directly from 95% ethanol to the
mounting medium, and covered with a cover slip. The
prepared slide may be viewed after 2 or 3 days, by which
time the radular teeth have stopped rotating. For best
results a compound microscope capable of a magnification
of at least 400 x should be used.
MATERIALS NEEDED
. saturated solution of trisodium phosphate
. concentrated solution of sodium hydroxide
. alcohol lamp, ringstand, ring and wire mesh
. 3 disposable hypodermic syringes
. 2very finely ground wooden-handled dissecting needles
. 70% ethanol
. 95% ethanol
. xylene
. canada balsam or other permanent mounting medium
10. eosin or eosin-type stain
11. 3 depression slides
12. 1 watch glass ( 1 - 2 inches in diameter)
13. 2 microscopes (1 dissection, 1 compound)
14. microscope slides and coverslips
ODUM TPwWDH we
Vol. 12; No. 1
THE VELIGER is open to original papers pertaining to any problem concerned
with mollusks.
This is meant to make facilities available for publication of original articles
from a wide field of endeavor. Papers dealing with anatomical, cytological, distri-
butional, ecological, histological, morphological, physiological, taxonomic, etc.,
aspects of marine, freshwater or terrestrial mollusks from any region, will be
considered. Even topics only indirectly concerned with mollusks may be acceptable.
It is the editorial policy to preserve the individualistic writing style of the
author; therefore any editorial changes in a manuscript will be submitted to the
author for his approval, before going to press.
Short articles containing descriptions of new species or other taxa will be given
preferential treatment in the speed of publication provided that arrangements
have been made by the author for depositing the holotype with a recognized
public Museum. Museum numbers of the type specimens must be included in the
manuscript. Type localities must be defined as accurately as possible, with geo-
graphical longitudes and latitudes added.
Short original papers, not exceeding 500 words, may be published in the column
“NOTES and NEWS’; in this column will also appear notices of meetings of
regional, national and international malacological organizations, such as A. M. U.,
U.M.E., W.S.M., etc., as well as news items which are deemed of interest to
our Members and subscribers in general. Articles on “METHODS and TECH-
NIQUES” will be considered for publication in another column, provided that
the information is complete and techniques and methods are capable of duplication
by anyone carefully following the description given. Such articles should be mainly
original and deal with collecting, preparing, maintaining, studying, photographing,
etc., of mollusks or other invertebrates. A third column, entitled “INFORMA-
TION DESK,” will contain articles dealing with any problem pertaining to
collecting, identifying, etc., in short, problems encountered by our readers. In
contrast to other contributions, articles in this column do not necessarily contain
new and original materials. Questions to the editor, which can be answered in this
column, are invited. The column “BOOKS, PERIODICALS, and PAMPHLETS”
will attempt to bring reviews of new publications to the attention of our readers.
Also, new timely articles may be listed by title only, if this is deemed expedient.
Manuscripts should be typed in final form on a high grade white paper, not
exceeding 81,” by 11”, at least double spaced and accompanied by a clear carbon
or photo copy. A pamphlet with detailed suggestions for preparing manuscripts
intended for publication in THE VELIGER is available to authors upon request.
A self-addressed envelope, sufficiently large to accommodate the pamphlet (which
measures 51,” by 8%”), with double first class postage, should be sent with the
request to the Editor.
EDITORIAL BOARD
Dr. Donatp P. Axszort, Professor of Biology
Hopkins Marine Station of Stanford University
Dr. Jerry DononveE, Professor of Chemistry
University of Pennsylvania, Philadelphia, and
Research Associate in the Allan Hancock Foundation
University of Southern California, Los Angeles
Dr. J. Wyatt Duruam, Professor of Paleontology
University of California, Berkeley
Dr. E. W. Facer, Professor of Biology
Scripps Institution of Oceanography, La Jolla
University of California at San Diego
Dr. Caner Hann, Professor of Zoology and
Director, Bodega Marine Laboratory :
University of California, Berkeley
Dr. G Datias Hanna, Curator
Department of Geology
California Academy of Sciences, San Francisco
Dr. Jorn W. Hepcretu, Resident Director
Marine Science Laboratory, Oregon State University
Newport, Oregon
Dr. Leo G. HERTLEIN,
Curator of Invertebrate Paleontology
California Academy of Sciences, San Francisco
EDITOR-IN-CHIEF
Dr. RupotF STouieR, Research Zoologist
University of California, Berkeley
: Scripps Institution of Oceanography, La Jolla
Dr. A. Myra KEEN, Professor of Paleontology and
Curator of Malacology
Stanford University, Stanford, California
Dr. Victor LoosanorrF, Professor of Marine Biology
Pacific Marine Station of the University of the Pacific
Dr: -“Joun McGowan, Associate Professor of
Oceanography
University. of California At. San Pies.
Dr. FRANK A. Pare praia af Yasloay
University of California; Berkeley
Mr. ALLyn G. Suir, Associate Giirator
Department of Invertebrate Zoology
California Academy of Sciences, San Francisco
Dr. Raupu I. Smiru, Professor of Zoology
University of California, Berkeley
Dr. Cuarzes R. Stasex, Associate Professor
of Zoology
Florida State University, Tallahassee, Florida
Dr. Donatp M. Wixson, Professor of Biology
Department of Biological Sciences
Stanford University, Stanford, California
ASSOCIATE EDITOR
Mrs. JEAN M. Cate
Los Angeles, California
S597¢,05
4.3
Astl.
THE
VELIGER
A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.
Berkeley, California
VOLUME 12 October 1, 1969 NuMBER 2
ContTENTS
Remarks on the Taxonomic Placement of Purpurellus JoussEAUME, 1880, with the
Description of a New Species (Gastropoda: Muricinae). (Plates 26, 27;
2 Text figures)
Witrmm K Emerson-« Antuony DiArmmio . . .. =... =.=... . 145
Two New Species of Galapagan Turrid Gastropods. (Plates 28, 29; 5 Text figures)
Wirram K> Emerson & Grorce EF. RapwiIN . ..... =... =. = - 149
Escape Responses of Three Indian Molluscs.
END) VANSELL ees SA esc, Semon C7)
Identification of the Beeding Ty pen in ate Genus Conus Iwacuet (Plate 30;
5 Text figures; 2 Tables)
C. EF Lim Bele : Ae ficwece yar TOO
Panamic Sites and rehveslowical roitesrs ae Dower Galicomiat (2 Tables)
LawreNcE H. FELDMAN. . . Bie) OS
Cyclostrema miranda BartscH, a Si ronyin GE Tatas hibcatinaeus (ioneden):
(3 Text figures)
Donatp R. Moore. . . . 169
Relative Growth Patterns of Two West Coast Squid ( Génatn fabric and Gonate ne
borealis). (2 Text figures)
Larry T. SPENCER . . . ely i
Occurrence of the Sacoglossan @pithonranch H ermaea dendntce fioents Hancore
in New England.
Kerry B. Crark & Davip R. FRANz . Soin dane aig
Observations on the Tentacles of Vaginulus boreliianus Covent (Mollusca: Gastro-
poda: Soleolifera) . (Plates 31 to 35; 3 Tables)
ARISTEO RENZONI . . Sh aon Loe rue ieee TO
Marine Fouling and Boring @reanignsn in Monterey Harbor — II. Second Year
of Investigation. (2 Text figures; 2 Tables)
Ei CBLUADERTIE Maan nist oc aut tly ec. MRO ache coe wie tte ase) ois ve va VOR
[Continued on Inside Front Cover]
Distributed free to Members of the California Malacozoological Society, Inc.
Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc.
Volume 12: $18. Domestic; $19.- in the Americas; 19.50 in all other Foreign Countries
Single copies this issue $9.00. Postage extra.
Send subscription orders to Mrs. JEAN M. CaTE, 12719 San Vicente Boulevard,
Los Angeles, California 90049. Address all other correspondence to Dr. R. STOHLER, Editor,
Department of Zoology, University of California, Berkeley, California 94720
Second Class Postage paid at Berkeley, California
VOM ax
I
ContENTs — Continued
Population Characteristics of Protothaca staminea (ConrapD) from Mugu Lagoon,
California. (9 Text figures)
Ronatp R. Scumipt & Joun E. WARME , . . Simeon
Mimicry of the Gastropod Mitrella carinata by the onpiipod Pleustes platypa.
(Plate 36)
Juves M. Crang, Jr. . . 200
Cypraeidae of the Red Sea at Maudie Eaiopia) nth a ‘Zooyeographical rAralyat’
based on the Scuitpers’ Regional Lists. (2 Text figures; 1 Map)
T.C. Fore L.P Russusn . . . . 201
The Distribution and Ecology of Sub- itera Species Gf Macoms (Bivalvia) off
Moresby Island and in Satellite Channel, near Victoria, British Columbia.
(9 Text figures; 3 Tables)
R. M. Dunnuit « D. V. Evttis . . . 207
Additional Bathymetric and Locality Data fone some lOnmthobranci nadl¢ an crept
from Santa Barbara County, California.
» . 193
Ricwarp S. Lez « Patrick BRoPpHY .. . BGS 6 CRO
A Bibliography of the Biological Writings of Eanes iSaomeann Gieeenmee!
EUGENE V. GOAN Gitte 0 os Ny ak ie eG Vetere CRP oe ten gles0 ke ten Mato 2 2.2
NOTES & NEWS (oii. ce05) ie) a inte 28) Se gout iste conus eo atirck ante) On RP oa 2
Russian Contributions to Malacology. KENNETH J. Boss
Report on some Abnormal Chitons from California and British Columbia.
GLENN BurcHarpT & Laura BURGHARDT
New Range for Mopalia hindsi recurvans BARNAWELL, 1960.
GLENN BurcHarRDT & LauRA BURGHARDT
Range Extension of Tylodina fungina in the Gulf of California.
James W. McBetu « R. Davin Bow.Lus
Ascophyllum nodosum: A Source of Exotic Invertebrates Introduced into
West Coast Near-Shore Marine Waters. RicHarp L. MILLER
Cadlina modesta: A Range Extension, with Notes on Habitat and a Color
Variation. (1 Text figure) Hans BertscH
A Note on the Opisthobranchs of Santa Cruz Island, California.
STEVEN J. Lonc
BOOKS, PERIODICALS & PAMPHLETS. .......... .. . 234
Note: The various taxa above species are indicated by the use of different type styles
as shown by the following examples, and by increasing indentation.
ORDER, Suborder, DIVISION, Subdivision, SECTION,
SUPERFAMILY, Famity, Subfamily, Genus, (Subgenus)
New Taxa
Vol. 12; No. 2
THE VELIGER
Page 145
Remarks on the Taxonomic Placement
of Purpurellus JOussEAUME, 1880,
with the Description of a New Species
(Gastropoda : Muricinae)
WILLIAM K. EMERSON
AND
ANTHONY D’ATTILIO
Department of Living Invertebrates, American Museum of Natural History
Seventy-ninth Street and Central Park West, New York, New York 10024
(Plates 26 and 27; 2 Text figures)
AMONG THE MOLLUSKS obtained by the “Ariel Expedi-
tion” to western Mexico in 1960 is a remarkable new
species of muricid gastropod. A single specimen was sorted
from a trawling sample obtained in the Gulf of California
by Dr. G. Bruce Campbell of Lynwood, California. Dr.
Campbell has kindly permitted us to describe it and he
generously deposited the holotype in the Los Angeles
County Museum of Natural History. A second specimen,
dredged in the Bay of Panama in 1935, was subsequently
found in the vast molluscan collections of the Allan Han-
cock Foundation. The gastropods of this collection have
been deposited on loan to the Los Angeles County Muse-
um of Natural History by the University of Southern Cali-
fornia. These specimens, both lacking soft parts, form the
basis for the description.
We take pleasure in naming this interesting discovery in
honor of Dr. James H. McLean, Curator of Mollusks, Los
Angeles County Museum of Natural History, who recog-
nized it in the Hancock Collection.
As a result of the present study, we have presented
anatomical evidence that requires the placement of Pur-
purellus JouSSEAUME, 1880, in the muricacid subfamily
Muricinae. Accordingly, we have re-assigned Purpurellus
from the Ocenebrinae to the status of a subgenus in the
muricid genus Pterynotus Swanson, 1833 (sensu lato).
MuRIGACEA
MourIciAgE
Muricinae
Pterynotus Swainson, 1833
Pterynotus Swainson, 1833, explanation to plate 100,
type species: Murex pinnatus Swainson, 1822 [=
Purpura alata Ropinc, 1798], by SD, Swarnson,
1833, explanation to plate 122. Voxes, 1964, plt. 14,
plt. 37, figs. 24 (shell), 51 (radula), 63 (operculum)
of Purpura alata.
Remarks: This genus has been traditionally placed in the
subfamily Muricinae owing to the presence of a typically
muricid shell and a muricoid operculum with a basal
nucleus. As pointed out by Voxes (1964, p. 15), the rad-
ular central tooth of the type species, Purpura alata, dif-
fers from that commonly found in the Muricinae in having
lost the smaller mtermediate cusps of the central tooth.
This condition may reflect a specialized feeding adaptation.
(Purpurellus) JoussEAUME, 1880
Purpurellus JouSSEAUME, 1880, p. 335, type species:
Murex gambiensis REEVE, 1845, by OD. VoxEs, 1964,
Page 146
p. 26, fig. 79 (shell of Murex gambiensis).
Triremis “Bayle MS” FiscHer, 1884, p. 641, type
species: Murex gambiensis Reeve, 1845, by OD.
THIELE, 1929, p. 288, fig. 313 (radula of Murex gam-
biensis).
Remarks: Vokes (1964, p. 26) recently afforded Purpur-
ellus JoUSSEAUME, 1880, subgeneric rank in the genus
Pteropurpura JoUSSEAUME, 1880, of the subfamily Triton-
aliinae [= Ocenebrinae]. This generic and subfamilial
placement was prompted largely by the similarity of the
shell morphology of the type species, Murex gambiensis
Reeve, 1845, with that of Pteropurpura (s.s.). She dis-
tinguished the shells of Purpurellus by their possession of
a greatly widened siphonal canal and the extreme inter-
ruption developed in the varical fringe between the body
whorl and the siphonal canal. Voxes noted that THIELE’s
(1929, p. 289) illustration of the radula of Murex gam-
biensis depicts muricine dentition and thus would require
placement of Purpurellus in the Muricinae, but she
questioned the authenticity of the figure and relied on
the apparent affinity of the shell morphology with that of
Pteropurpura for subfamilial assignment.
We have found, however, the radular dentition of
Murex pinniger Broverip, 1833, an eastern Pacific repre-
sentative of Purpurellus, to be muricine (Text figure 1)
and to be reminiscent of THIELE’s radular figure of M.
gambiensis (Text figure 2). Both species are now deter-
Figure 1
Pterynotus (Purpurellus) pinniger (BRoveErIP, 1833)
Rachidian and lateral dentition, approximately 300; specimen
obtained by fishermen out of Guaymas, Sonora, Mexico;
L. Thomas collection
THE VELIGER
Vol. 12; No. 2
mined to have muricine radulae, necessitating placement
of Purpurellus in the subfamily Muricinae. We, therefore,
have returned Purpurellus to the status of a subgenus in
the genus Pterynotus (sensu lato), where it previously
was assigned (Emerson, 1960).
Figure 2
Pterynotus (Purpurellus) gambiensis (REEVE, 1845)
Rachidian and lateral dentition, greatly enlarged
{after THIELE, 1929, p. 289, fig. 313]
The following Recent species are known to be referable
to Purpurellus:
1. Murex gambiensis Reeve, 1845 [= ? Murex osseus
REEVE, 1845], the type species, from the tropical eastern
Atlantic.
2. Murex pinniger Broverip, 1833 [= Centrifuga inezana
Duruam, 1950, fide SHaAsky & CAMPBELL, 1964, p. 116],
from the tropical eastern Pacific.
3. Pterynotus (Purpurellus) macleani, spec. novy., described
herein, from the tropical eastern Pacific.
In addition to the three living species, extinct species of
Purpurellus are known from the Early Miocene of North
Carolina, the Miocene of France, and the Pliocene of
Italy (VoKEs, in litt.). The available distributional data,
therefore, indicate that the living representatives of this
group are surviving, relict elements of the older Tertiary,
west-Tethyan faunas. This would explain the present ap-
parently discontinuous distribution, with species now living
in the tropical eastern Atlantic and the eastern Pacific
Oceans, but not occurring in the western Atlantic.
The status of Centrifuga inezana DurHAM, 1950, pp.
113, 114, plt. 26, figs. 1, 4 (here illustrated, Plate 27,
Explanation of Plate 26
Pterynotus (Purpurellus) macleani EMERSON & D’ATTILIO, spec. nov.
Figures 1, 2: Holotype, Loreto Channel, Baja California, Mexico,
in 25 fathoms; 2. (Note open siphonal canal and immature
outer lip).
Figures 3, 4: Paratype, off Secas Island, Panama, in 12 fathoms;
2. (Note closed siphonal canal and mature stage of apertural
lip) .
Pterynotus (Purpurellus) pinniger (BRODERIP, 1833)
Figures 5, 6: Bicolored juvenile specimen of 5 postnuclear whorls,
Santa Rosalia, Isla San Marcos, Baja Califomia, Mexico,
Ben and Ruth Purdy collection; <2.
Figure 7: Juvenile specimen of 6+ postnuclear whorls, Panama
Bay, Panama, dredged, Lee and Helen Beils leg.,
ex- Ruth Craine collection; X2.
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Tue VELIGER, Vol. 12, No. 2
Figure 7
Figure 6
Figure 5
Wol 2 Now 2
THE VELIGER
Page 147
Figures 1, 2) requires additional comment. On the basis
of the limited data previously available (EMERSon, 1960,
pp. 10-12, figs. 6, 7), we concluded that DurHAm’s
taxon was referable to the subgenus Purpurellus, but we
could not determine with certainty its conspecificity with
Pterynotus(Purpurellus) pinniger (BRopERIP, 1833). VoKES
(1964, p. 26) subsequently pointed out that the type
localities of these taxa (Ecuador for pinniger and the
Gulf of California for inezana) were geographically sep-
arated by some 3000 miles. SHasky & CAMPBELL (1964,
p. 116) reported on additional specimens and they con-
cluded that the two nominal species were synonymous.
One of Shasky’s specimens from the Guaymas area in the
Gulf of California is illustrated herein (Plate 27, Figures
3, 5), and it can be assigned with confidence to P (P)
pinniger, cf. Plate 27, Figures 5, 6. Although the holotype
of Pterynotus (Purpurellus) inezana (DURHAM, 1950) is a
poorly preserved (broken and water worn) Pleistocene
specimen, the presence of P (P) pinniger in the Gulf of
California serves to substantiate the findings of SHasky
& CAMPBELL, and we have placed znezana in the synonymy
of pinniger.
Pterynotus (Purpurellus) macleant EMERSON & D’ATTILIO,
spec. nov.
(Plate 26, Figures 1 to 4)
Diagnosis: A small muricid of 7 postnuclear whorls with
broadly expanded varices. The shell is similar to that of
Pterynotus (Purpurellus) pinniger, but the much smaller
shell has a stouter appearance. Specimens of the new spe-
cies with 7 postnuclear whorls are less than } the size of
specimens of this congenitor with the same number of
whorls.
Description: Shell is small, translucent pink, polished, of
thin and fragile appearance, with three wing-like varices;
the 7 whorls blend into a protoconch of 14 convex whorls.
Spire is slightly convex, with a well defined suture and
on the later whorls has a tubercle-like bulge which be-
comes reduced near the sutural groove. The body whorl is
swollen in the area of the large tubercle at the shoulder.
The three thin blade-like varices have a sinuous contour;
they overlap from one whorl to the next forming, in part,
on the spire, one continuous blade, demarked only by
sutural indentations. On each whorl the varices, just above
their broadest part, extend into upturned triangular
spines. The edge of the varices is crenulated and wavy.
Viewed from the aperture side, the varices on each whorl
have an overlapping, folded-over layer which is tightly
appressed; the edge of the overlapping layer begins at
the tip of the triangular spine and curves down in a
gradually widening segment.
The aperture is ovate, small and entire; it is notched
slightly above the opening to the siphonal canal, and in
addition it is notched where the double layered portion
of the varical spine begins. Both parietal and labial areas
of the apertural lip are erect. The siphonal canal is long,
broad, and is slightly opened in the holotype and closed in
the paratype. The edge of the canal, on the parietal side,
adherently overlies along its length a portion of the canal;
the canal is not recurved except for the tube-like open
distal end which is sharply bent back. Two previous canals
remain on the siphonal fasciole. The shell is microscop-
ically, spirally striate except for some strong striae present
in the region of the larger body tubercle.
Color: In the holotype, there are faint red-brown spiral
bands at the aperture on the edge of the labrum, with 5
additional spiral bands on the canal. The spiral color
bands rapidly fade away from their strong beginning. They
persist, however, on the attachment portions of earlier
varices. The paratype lacks coloration (owing to fading?)
and is porcellaneous white.
Opercular and radular characters are not known.
Holotype: Los Angeles County Museum No. 1230, trawled
by the “Ariel Expedition,’ in 25 fathoms from Loreto
Channel, Baja California, Mexico, on gravel and coral
shell bottom, August 29, 1960 (type locality). Sorted from
trawl samples by G. Bruce Campbell. Length 29.0 mm,
width 21.9 mm.
Paratype: Los Angeles County Museum No. A.H.E
448-35, dredged by the Hancock Foundation vessel
“Velero III” in 12 fathoms in sand off Secas Island, Pan-
ama, 7°57'45”N, 82°00’45” W, February 5, 1935
(Fraser, 1943). Length 22.4 mm, width 12.7 mm.
Remarks: The shell of the new species is morphologically
similar to that of Pterynotus (Purpurellus) pinniger (Brop-
ERIP) of the eastern Pacific and P (P) gambiensis (REEVE)
from western Africa. It differs from the latter two species
by its much smaller size; its comparatively greater width
relative to height; and the lack of an indentation in the
varical flange between the body whorl and the canal. The
lamella in both P (P) gambiensis and P. (P) pinniger tend
to have their edges more crenulated and recurved into
spiny points and the spiral cords more developed. Ptery-
notus (P.) pinniger has a larger, heavier, more elongate
shell. All three of the Recent species have a similar pattern
of coloration; the spiral color bands start somewhat
strongly at the labrum and fade out rapidly before reach-
ing the next varix. The lamellae on all three species are
continuous with the body of the shell on their dorsal side.
On their ventral side there is a distinct break between
the thickened varices and the lamellae extending above;
Page 148
THE VELIGER
Vol. 12; No. 2
in addition, the sculpture of the ventral sides of the lamel-
lae is finely multilaminate.
The opercula of Pterynotus (Purpurellus) pinniger and
P (P) gambiensis have a nucleus somewhat below and to
the right of center.
ACKNOWLEDGMENTS
In addition to Drs. G. B. Campbell and J. H. McLean,
and E. H. Vokes, we are most grateful to the following
friends and colleagues for the loan of pertinent specimens:
Mr. and Mrs. John Q. Burch of Seal Beach, California,
Mrs. Ruth A. Craine of Norwich, New York, Dr. Donald
R, Shasky of Redlands, California, Mr. Lawrence Thomas
of Morro Bay, California, Mr. Joseph H. Peck, Jr., Muse-
um of Paleontology, University of California, Berkeley,
and Mr. and Mrs. Ben Purdy of San Diego, California.
Dr. George E. Radwin of the San Diego Natural History
Museum kindly provided a radular mount of Murex pin-
niger from which the present illustration was drawn. Our
colleagues, Miss Lynne Judge and Mr. William E. Old,
Jr., assisted us in various ways.
LITERATURE CITED
BropeErip, WILLIAM JOHN
1833. | Characters of new species of Mollusca and Conchifera
collected by Mr. Cuming. Proc. Zool. Soc. London for
1832: 173 - 179 (14 January 1833)
DuruaM, Joun Wyatt
1950. 1940 E. W. Scripps cruise to the Gulf of California. Pt.
II. Megascopic paleontology and marine stratigraphy.
Mem. Geol. Soc. Amer., no. 43: 1 - 216; 48 plts.
EMERSON, WILLIAM KEITH
1960. | Remarks on some eastern Pacific muricid gastropods.
Amer. Mus. Novitates no. 2009: 15 pp.; 7 figs
FiscHer, PAuL
1884 [1880 - 1887]. Manuel de conchyliologie et paléontologie
conchyliologique. Paris, 1369 pp.; 23 plts. (fasc. 7, Muri-
cidae, publ. in 1884)
Fraser, CHARLES McLEAN
1943. General account of the scientific work of the Velero III
in the eastern Pacific, 1931-41. Part III. A ten-year list of
the Velero III collecting stations (charts 1 - 115). Allan
Hancock Pacif. Exped. 1 (3): 256-431; 115 charts. Univ. So.
Calif. Press, Los Angeles, California
HAN Ley, SYLVANUS CHARLES
1841. Exotic conchology, by Witttam Swainson. Second ed.
London, 39 pp.; 48 plts.
JoussEAUME, FELIX PIERRE
1880. Division méthodique de la famille des purpuridés,
Le Naturaliste, Paris, 1 (42): 335 - 336
REEvE, LovELL Aucustus
1845-1846. Conchologia Iconica: or Illustrations of the shells of
molluscous animals. Monograph of the genus Murex.
3: plts. 1 - 36; 1 suppl. plt. (April 1845 to April 1846)
Ropinc, PETER FriepRIcH
1798. Museum Boltenianum ... : pars secunda continens con-
chylia sive testacea univalvia, bivalvia & multivalvia.
Hamburg (Johan Christi Trappii) pp. i- viiit+1- 199
SHASKY, DONALD R. & G. BRUCE CAMPBELL
1964. New and otherwise interesting species of mollusks from
Guaymas, Sonora, Mexico. The Veliger 7 (2): 114-120;
plts. 21, 22: 1 text fig.; 1 map (1 October 1964)
Swainson, WILLIAM
1822. Descriptions of several new shells, and remarks on others,
contained in the collection of the late Mrs. Bligh,
as appendix zn C. Dusots, A catalogue of the rare and valuable
shells which formed the celebrated collection of the late Mrs.
Bligh, London, illus. [not seen; Swatnson’s Bligh appendix
reprinted by SytvaNus Han -ey, 1841].
1829 - 1833. Zoological illustrations or original figures and
descriptions of new, rare, or interesting animals, selected
chiefly from the classes of ornithology, entomology, and conch-
ology, and arranged according to their natural affinities.
London, (Baldwin « Cradock), ser. 2, vol. 3, The shells.
THIELE, JOHANNES
1929[-1931]. Handbuch der systematischen Weichtierkunde.
Jena, Gustav Fischer, 1929 - 1935; 1154 pp.; 893 text figs. (pp.
1-376 publ. in 1929)
VoKEs, Emity H.
1964. | Supraspecific groups in the subfamilies Muricinae and
Tritonaliinae (Gastropoda : Muricidae).
(1): 1-41; plts. 1-3; text figs.
Malacoiscia
Explanation of Plate 27
Pterynotus (Purpurellus) pinniger (BRODERIP, 1833)
Figures 1, 2: Holotype of Centrifuga inezana DurHAM, 1950, pl. 26,
figs. 1, 4, specimen with 4+ postnuclear whorls (early whorls
lost), Pleistocene, Punta Santa Inez, Baja California, Mexico;
approximately X2.
Figures 3, 5: Specimen of 6+ postnuclear whorls, off Cabo Haro,
Guaymas, Sonora, Mexico, 20 to 40 fathoms.
Donald Shasky collection; <2.
Figure 4: Lectotype of Murex pinniger (BRopERIP, 1833), Xipi-
xapi, Ecuador, 8 fathoms, after EMERSON, 1960, fig. 7; X2.
Figure 6: Specimen, partly encrusted with bryozoa, with 5+
postnuclear whorls, off Cabo Haro, Guaymas, Sonora, Mexico,
30 fathoms, G. Bruce Campbell collection; 2.
[EMERSON & D’ArTILI0] Plate 27
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THE VELIGER, Vol. 12, No. 2
Wolel2 No; 2
THE VELIGER
pagesta9
Two New Species of Galapagan Turrid Gastropods
WILLIAM K. EMERSON
Department of Living Invertebrates, American Museum of Natural History
Seventy-ninth Street and Central Park West, New York, New York 10024
AND
GEORGE E. RADWIN
Natural History Museum, Balboa Park, P. O. Box 1390, San Diego, California 92112
(Plates 28 and 29; 5 Text figures)
AMONG NUMEROUS MARINE MOLLUSKS kindly submitted
by Mrs. Jacqueline DeRoy of Academy Bay, Isla Santa
Cruz, the Galapagos Islands to the senior author for
study are representatives of two new species of toxoglos-
sate gastropods referable to the Turridae. The description
of these interesting discoveries forms the basis of this
report.
We take great pleasure in dedicating this paper to our
colleagues, Dr. Leo G. Hertlein of the California Aca-
demy of Sciences and Dr. A. Myra Keen of Stanford
University, both of whom have made numerous contribu-
tions to Malacology.
Hindsiclava HERTLEIN & STRONG, 1955
Hindsiclava HERTLEIN & STRONG, 1955, p. 227; type
species: Pleurotoma militaris “Hinps” Reeve, 1843,
by OD.
Turrigemma Berry, 1958, p. 88, type species: Turri-
gemma torquifer Berry, 1958, by SD (Berry, 1964,
154). Otsson (1964, p. 96) referred BeRRy’s Tur-
rigemma to the synonymy of Hindsiclava.
Remarks: The genus Hindsiclava was proposed for shells
possessing strong axial ribs and cords to give a reticulate
sculpture to the surface and with a well-developed node
of callus at the sutural edge of the anal sinus. This anal
node is apparently a gerontic character that does not
appear in specimens with immature outer lips or in
juveniles.
The following nominal species from the Eastern Pacific
are apparently referable to this genus, in addition to the
type species, Pleurotoma militaris REEVE, 1843.
1. Turris (Surcula) resina Dati, 1908, p. 264, Gulf of Panama, in
322 fathoms (U.S. Fish Commission Station 3345), based on a
single, large incomplete specimen with an anal node; Dati, 1919,
p. 16, plt. 2, fig. 4 (holotype). This taxon was placed in the syn-
onymy of Hindsiclava militaris by Otsson, 1964, p. 96.
2. Turris (Surcula) dotella Dati, 1908, p. 263, off Baja California,
Mexico, 27 fathoms, holotype (U.S. Fish Commission Station
2823) and 2 paratypes off La Paz, Baja California, Mexico;
Keen, 1958, p. 464, fig. 842 (holotype, U.S. N.M. No. 96731),
an immature specimen lacking an anal node.
3. Turris (Surcula) notilla Dai, 1908, p. 263, off Cape San Lobos,
Baja California, Mexico, in 58 fathoms, holotype (U.S. Fish
Commission Station 3017), and 2 paratypes from same station.
KEEN, 1958, p. 476, fig. 906 (holotype U.S. N.M. No. 110602),
an immature specimen lacking an anal node, strongly resembling
Hindsiclava dotella but with stronger axial sculpture.
4. Clathrodrillia andromeda Dauu, 1919, p. 16, plt. 2, fig. 2, off La
Paz, Baja California, Mexico, in 10 fathoms; Kren, 1958, p.
464, fig. 841 (copy of Datt’s (1919) figure). The holotype is
a small juvenile specimen, lacking an anal node.
5. Turrigemma torquifer Berry, 1958, p. 88, off Puerto Refugio,
Angel de la Guarda Island, Mexico, in 65 fathoms. KEEn, 1958,
p. 465, fig. 845 (holotype in collection of Dr. S.S. Berry?). This
nominal species appears to be a junior synonym of Hindsiclava
andromeda (Datt, 1919) ; the figured specimen of BErrRy’s taxon
apparently represents a mature specimen.
6. Hindsiclava hertleini, new species, described below, from the
Galapagos Islands.
The genus Hindsiclava is apparently limited in fossil
and present day distribution to the New World. In addi-
tion to the record of Hindsiclava militaris REEVE in the
late Neogene of Ecuador (Otsson, 1964), several nominal
Page 150
species that appear to be referable to this genus are known
from numerous deposits of Mio-Pliocene age in the Carib-
bean region. For example, “Pleurotoma” consors SowER-
BY (1850, p. 50), originally described from the Miocene
of the Dominican Republic, was stated by SoweRBy to
be an analogue of REEvE’s militaris. RutscH (1934, pp.
99 - 101, figs. 13 - 16) cited specimens of SoweErRBy’s spe-
cics from the Neogene of Venezuela and he discussed
representatives of the group of Hindsiclava consors (Sow-
ERBY) then known from the East American region. He
also pointed out the similarity of “Drilla” alesidota DALL
(1889, p. 84) and “Drilla” macilenta DALL, (1889, p. 85,
plt. 36, fig. 1), Recent species described from the Gulf of
Mexico and off Barbados, to Hindsiclava consors.
Hindsiclava militaris REEVE, 1843, ex Hinps MS
(Plate 28, Figures 1 to 4; Text figures 1, 2)
Pleurotoma militaris REEVE, 1843 [March], vol. 1, Pleurotoma,
sp. 55, plt. 7, fig. 55, “Veragua, Central America’’[Pana-
ma], 18 fathoms, mud, Hinds [type locality].
Clavatula militaris Hinps, 1843 [July], p. 38, Veragua [Pana-
ma], 18 fathoms, Panama, 8 - 30 fathoms. Hinps, 1844,
p. 16, plt. 5, fig. 10, Veragua, Central America [Panama],
in 18 fathoms; Panama, in 8 - 30 fathoms.
Drillia militaris “Hinps”, Tryon, 1884, vol. 6, p. 181, plt.
10, fig. 74 (copy of REEve’s (1843) figure).
Pleurotoma (Drillia) militaris REEVE, WEINKAUFF, 1887, vol.
4, div. 3, p. 132, plt. 29, fig. 10.
[?] Turris (Surcula) resina Dat, 1908, p. 264, Gulf of Pana-
ma, in 322 fathoms, mud, U.S. Fish Commission ‘“Alba-
tross” station 3354, 7°09’45” N, 80°50’00” W [type lo-
cality].
[?] 2Clathrodrillia resina Dat, Dat, 1919, p. 16, plt. 2, fig.
4 (holotype; U.S.N.M.No. 123103); Pmssry & Ots-
son, 1941, p. 18, plt. 2, fig. 1, Punta Blanca, Ecuador,
Canoa formation, Pliocene.
Hindsiclava militans “Hinps”, HERTLEIN « STRONG, 1955, p.
227, [in part, excluding reference to Clathrodrillia aeno-
ne Dati, 1919], Pinas Bay, Panama, 14 - 33 meters; Ardita
Bay, Colombia, 34-43 meters; Octavia Bay, Colombia,
24-28 meters, in gray sand, sandy mud, and _ black
bottom. Orsson, 1964, p. 97, plt. 17, figs. 3, 3a, Quebra-
da Camerones, Ecuador, Esmeraldas formation, late Mio-
cene-early Pliocene. KEEN, 1958, p. 465, figured on
frontispiece.
Range: Off Magdalena Bay, Baja California, Mexico,
through the Gulf of California, and south to Octavia Bay,
Colombia (HERTLEIN & STRONG, 1955), off shore to 90
fathoms, ? 322 fathoms. Also reported as a fossil from late
Neogene deposits of Ecuador (Pitssry « Oxsson, 1941;
Otsson, 1964). ;
Radular characteristics: Radular formula 1-0°0-0-1.
A racemose cluster of 30-40 “duplex” uncinal teeth
THE VELIGER
Vol. 12; No. 2
attached to an almost invisible, vestigial radular ribbon
and seemingly enclosed in a membranous sac (Text figure
1). Each tooth consists of two articulating parts, a broad,
blade-like member and a narrower, stylet-like member.
The two parts articulate proximally and lie parallel for
most of their length (Text figure 2). The radular arrange-
Figure 1 Figure 2
Hindsiclava militaris (REEVE, 1843)
Figure 1: Several radular teeth in their natural grouping.
Figure 2: Single tooth; both figures greatly enlarged.
ment in Hindsiclava militaris, the type species of Hindsi-
clava, suggests placement of this genus in the turrid sub-
family Crassispirinae (see Morrison, 1966).
Remarks: The type locality of Hindsiclava militaris is off
Veragua, Panama, in 18 fathoms, based on specimens
collected on the world-wide voyage of “H. M.S. Sulphur,”
under the command of Captain Belcher, in 1836 - 1842
(see Hinps, 1844, p. 16, plt. 5, fig. 10). Hrnps’ preliminary
description of this taxon did not appear until October of
1843 (Hinps, 1843), whereas REEveE’s description and
figure of the taxon based on Hinp’s manuscript name ap-
peared in March of 1843 (Reeve, 1843). Therefore, the
authorship of this species must be credited to REEVE, 1843,
on the basis of priority, as there is no evidence that H1nps
provided the description that appeared in REEve’s (1843)
monograph. It also should be noted that REEvE’s descrip-
tion and those of Hinps (1843, 1844) are not identical.
This fact further suggests that REEve (1843) prepared his
own description for this taxon.
KEEN (1966, p. 272) was unable to locate the type spe-
cimens of Pleurotoma militaris REEvE in the Belcher-
Hinds collection of the British Museum (Natural History).
Before undertaking the description of the new species of
Hindsiclava that appears below, we inquired of the offi-
cials of the Mollusca Section of the British Museum
Vol. 12; No. 2
regarding the presence of potential type material of
REEVE’s taxon in their collection. They informed us that
documented type specimens could not be located, but
they kindly provided us with photographs of 3 specimens
labeled “Pleurotoma militaris Hinps, Central America”
from the Hugh Cuming Collection. Although these speci-
mens do not now possess definite locality data, they may
represent part of the original lot.
In the absence of better documented material, these
specimens are candidates for syntypic status. They are
worn specimens, the largest two being slightly smaller
(measuring 39 mm and 36 mm, respectively, in length),
than the specimen illustrated by REEveE (1843, plt. 7, fig.
55), which, judging from the figure, is 45 mm in length.
The largest and best preserved specimen from the Cuming
Collection is illustrated here, Plate 28, Figure 2, for
comparison with a copy of REEve’s original illustration,
here copied, Plate 28, Figure 1."
An examination of 12 lots of Hindsiclava miltaris in
the collection of the American Museum, totaling nearly
75 specimens, reveals that a well-developed anal node is
present on only 5 of the larger specimens. The smallest
specimen possessing a node measures 42.5 mm in length,
whereas the largest with a node measures 45.6mm in
length. Similar-size specimens with immature outer lips
lack a node (cf. Plate 28, Figures 3 and 4). Larger speci-
mens attaining a maximum length of 50 mm, all of which
appear to have immature outer lips, lack nodes. These
data suggest that the presence of an anal node is a gerontic
character of genetic significance that does not occur in
juvenile individuals or in old specimens that have not
developed a mature outer lip. It is possible, however, that
the development of the node is the result of sexual di-
morphism, or represents some other biological factor,
but such conclusions must await further study.
We have questionably placed Hindsiclava resina (DALL,
1908) in the synonymy of ReEEve’s H. militaris. DALL’s
taxon was based on a single, incomplete specimen, that
measures approximately 50 mm in length. This specimen
possesses a partially developed anal node (Dati, 1919,
plt. 2, fig. 4). It was stated to have been dredged in 322
fathoms in the Gulf of Panama. All live-collected speci-
mens of Hindsiclava militaris that we have seen are from
' According to Dance (1966, p. 213), “The British Museum
(Natural History) purchased 277 shells from [G. B.] Sowerby
[3'4 of name], many of them described and figured in the Mol-
lusca reports of the Sulphur and Samarang voyages.” Therefore,
it would appear that part of Hind’s Sulphur-material was dis-
tributed by the Sowerbys who were shell dealers, and the present
specimens might have been obtained by Hugh Cuming from this
source, or they could have been given to Cuming by Belcher or
Hinds (see KEEN, 1966).
THE VELIGER
Page 151
stations on the continental shelf. Possibly this specimen
was carried by turbidity currents or by some other means
down the slope to this depth. Orsson (1964, p. 96)
placed H. resina in the synonymy of H. militaris, and
he stated: “Dati’s Clathrodrillia resina was described
from a broken specimen (its length when perfect esti-
mated at 88 mm.) dredged from a depth of 322 fathoms
in the Bay of Panama.” We have not seen specimens of H.
militaris that approach this size. HERTLEIN & STRONG
(1955, p. 227), however, cite “A very large specimen [of
H. militaris] in the collections of the California Academy
of Science from the southern portion of the Gulf of Cali-
fornia: length, 56.4 mm.; maximum diameter, 1.7 mm.”
Da. (1908, p. 264) also mentions with reference to
the description of Hindsiclava resina: “A large shell very
much broken and eroded with a somewhat similar form
and sculpture ” was dredged from 134 fathoms,
near Cocos Island. The incomplete specimen measures
about 58 mm in length and “It has a large amorphous
mass of callus on the proximal end of the pillar,
and [this] may be a pathological feature.”
This would appear to be a reference to a well-developed
anal callus. Dati did not propose a name for the Cocos
Island specimen because of the incompleteness of the shell.
Clathrodrillia aenone Datu (1919, pp. 15 - 16), which
was not illustrated at the time the taxon was proposed,
was referred to Hindsiclava by HERTLEIN & STRONG
(1955, p. 227). The holotype of Datt’s taxon was sub-
sequently illustrated by Kren (1958, p. 454, fig. 770),
who retained it in Clathrodrillia.
vt}
Hindsiclava hertleint EMERSON & RaADWIN, spec. nov.
(Plate 28, Figures 5, 6; Text figure 3)
Diagnosis: A medium-sized, slender species of Hindsicla-
va, characterized by numerous, coarsely nodulose, axial
ribs (23 on body whorl of holotype), crossed by dark
chestnut-brown lirations, over an of-white base. Aperture
white, tinged with buffyellow.
Description: Shell slender, spire acuminated, whorls
shouldered; axial sculpture of slightly nodulose ribs, 20 to
23 on the body whorl; spiral sculpture of depressed lirae
which decussate the axial ribs to form coarse nodules;
body whorl with 7 major spirals of equal size and 1
minor spiral just below the suture, with 5 minor spirals at
the base of the siphonal canal; spire consists of 33 smooth
nuclear whorls, and 8 sculptured, postnuclear whorls,
which become progressively more strongly ornamented
posteriorly; a narrow, weakly nodulose sutural collar is
present above the suture on the spire, followed by 2
major nodulose axials, each divided by a narrow depressed
Page 152
lira; outer lip thin with a moderately developed anal
notch, lacking an anal callus in the holotype and the
paratypes (presumably because of the immaturity of the
specimens). Exterior off-white, with spirals and sutures
dark chestnut-brown; interior white with inner lip and
parietal wall tinged bright buff-yellow. Periostracum thin,
nearly transparent, pale straw color.
Radular characteristics: Radular formula 1:0:0-0-1.
A racemose cluster of approximately 30 “duplex” uncinal
teeth attached to a somewhat vestigial radular ribbon
and seemingly enclosed in a membranous sac. Each
tooth consists of two parts, a broad, blade-like member
and a narrower, stylet-like member. The two parts articu-
late proximally and lie parallel for most of their length
(Text figure 3). This type of radular dentition, typical of
Hindsiclava, as shown earlier, suggests that this species
and H. militaris should properly be assigned to the turrid
subfamily Crassispirinae (see Morrison, 1966).
Measurements: holotype 36.5 mm in length, 11.6 mm in
width; largest paratype 31.7:mm in length, 9.1 mm in width
(incomplete specimen).
Figure 3
Hindsiclava hertleini EMERSON & RADWIN, spec. nov.
Single radular tooth; greatly enlarged.
THE VELIGER
Volel2 Now
Type locality: South of Isla Daphne, Galapagos Islands,
in 101 meters. J. DeRoy, November 25, 1967 (holotype,
see Plate 28, Figure 5; and 1 paratype).
Type depositories: holotype, A. M.N.H.No. 150514;
1 paratype A.M.N.H.No. 152603 (dead specimen),
May 27, 1967, Plate 28, Figure 6); 1 paratype S.D.N.
H. S. No. 50768, December 9, 1968; 1 paratype, A. D’At-
tilio Collection, December 9, 1968; all from off Santa Cruz
Island, Galapagos Islands in 73 - 91 meters.
Remarks: The holotype is a fresh, apparently live-taken
specimen, whereas one paratype is a dead-collected,
bleached specimen lacking the early whorls of the spire
and part of the outer lip. The two other paratypes are
live-collected specimens. All specimens appear to have
immature outer lips and lack an anal callus in this stage
of development.
The new species resembles Hindsiclava militaris in
general features but differs in the narrower, sutural collar,
more angular shoulder, fewer spirals and coarser nodes on
the body whorls, as pointed out by Dr. Hertlein (in litt.),
who kindly examined two of the Galapagan specimens.
The periostracum of H. militaris is a greenish-brown
while that of the new species is a pale straw-yellow.
Lioglyphostoma variculosa (SowERBy, 1834, p. 139;
Reeve, 1843, plt. 22, fig. 194), described from off west
Panama and reported also from Mazatlan, Mexico (KEEN,
1958, p. 464), appears to have similar ornamentation, but
it has fewer axial ribs. We have not seen specimens re-
ferable to this taxon.
Mitrolumna Bucqguoy, DAUTZENBERG
« DoLiLFus, 1883
Mitrolumna Bucguoy, DauTzENBERG & Do.irus, 1883, p.
121; type species: Mitra olivoidea CANTRAINE, 1835, by
OD. PowELL, 1966, p. 67 (synonymy and remarks).
Clinomitra BELLARDI, 1889, p. 152; type species: Clinomitra
rovasendae Breviarpi, 1889, by M, a Miocene fossil.
Diptychomitra BELLARDI, 1889, p. 152; type species: Diptycho-
mitra eximia BELiarpI, 1889, by SD (Cossmann, 1899,
p. 174), a Miocene fossil.
Explanation of Plate 28
Figures 1, 2a, 2b, 3a, 3b, 4a to 4c: Hindsiclava muilitaris REEVE,
1843, type species of Hindsiclava HERTLEIN & STRONG, 1955.
Figure 1: copy of illustration of Hinps, 1843, plt. 5, fig. 10; 1.25.
Figures 2a, 2b: probable syntypes, Hugh Cuming Collection, British
Museum (photograph courtesy and © of British Museum (Natural
History) ); 1.25.
Figures 3a, 3b: 146-165 meters, Gorda Banks, Baja California,
Mexico, Templeton Crocker Station 150-D-12; 1.25 (note well-
developed anal node).
Figures 4a to 4c: 101 - 183 fathoms, Arena Bank, Baja California,
Mexico, Templeton Crocker Station 136-D-17; X1.25 (note the
lack of an anal node).
Figures 5a, 5b, 6a, 6b: Hindsiclava hertleini EMERSON & RADWIN,
spec. nov. Figures 5a, 5b: Holotype, with periostracum; X2.
Figures 6a,6b: Paratype, dead, worn specimen, lacking early whorls;
X2.
THE VELIGER, Vol. 12, No. 2 [EMERSON & Rapwin] Plate 28
He
E
E
E
=
E
E
Figure 3a Figure 3b Figure 4 a Figure 4 b Figure 4 c
Figure 6 a Figure 6 b
Vol. 12; No. 2
THE VELIGER
Page 153
Remarks: Originally established for species having shells
intermediate in appearance between some species of Mitra
and Columbella, this genus appears to be related closely
to Mitromorpha CarPENTER, 1865 (not Mitromorpha A.
ApaMS, 1865), type species: Mitromorpha filosa CARPEN-
TER, 1865 [= M. carpenteri GuiBEert, 1954]. The major
difference seems to be the presence of two well-formed
median pillar plicae in Mitrolumna. The radular dentition
suggests a close relationship between these two genera,
cf. Text figures 4, 5 herein with the text figure 123 of
Powe Lv (1966), a radular illustration of the type species
of Mitromorpha.
Until the discovery of the new Galapagan species, the
only living representative of the genus Mitrolumna known
was the type species M. olivoidea CANTRAINE, 1835 from
the Mediterranean and north Atlantic. In addition, the
following Miocene species were assigned to Mitrolumna
by PowE.y (1966, p. 67): canaliculata BELLARDI, 1889;
clathrata BELLARDI, 1889; cancellata PEyYRoT, 1938; doll-
fusi Pryrot, 1938; filifera BELLARDI, 1889; michaudi
MicHeE.Lottl, 1847; and rovasendae BELLARDI, 1889. Mit-
rolumna olivoidea is also known from the European Plio-
cene and Pleistocene.
It should be noted in passing that the well-known spe-
cific name filosa CARPENTER in the genus Mitromorpha
CarRPENTER was validly replaced with M. carpenteri by
Gupert (1954, p. 43) owing to the fact that GLIBERT
had assigned a European Miocene species, Columbella
filosa Dujarpin (1837, p. 302), to the genus Mitromor-
pha (s. s.). Thus, CARPENTER’s specific name filosa became
a junior secondary homonym, and Mitromorpha carpen-
tert was proposed by GLiBERT as a replacement name for
M. filosa CarPENTER, 1865, not M. filosa (Duyjaronin,
1837).
Mitrolumna olwoidea (CANTRAINE, 1835)
Mitra olivoidea CANTRAINE, 1835, p. 391, Mediterranean, Ad-
riatic. TOMLIN & SHACKLEFORD, 1914, p. 245, Sao Thome,
Canary Islands, and Mediterranean.
Mitra columbellaria Scaccut, 1836, p. 10, figs. 12, 13, Naples,
Italy.
Mitra obsoleta Puiipr1, 1836, (not “Broccut,’ GRATELOUP,
1834, p. 290), vol. I, p. 230, Sicily, Italy.
Columbella greci Putri, 1844, vol. II, p. 194, plt. 27, fig.
18, Sicily, Italy.
Mitra striarella Catcara, 1841, p. 66, Sicily, Italy.
Mitra clandestina ReEve, 1845, vol. II, sp. 253, plt. 32, fig.
263, no locality.
Mitra columbellaris Scaccui, Petit, 1860, vol. 8, p. 258, “les
cétes de la Provence,” (France).
Mitrolumna olivoidea Bucquoy, DauTzENBERG & DOLLFUS,
18835 pan lee pli lon tess 33 .- Som (type), 301-139
(‘‘var.” [ieties]), Roussillon, France; synonymy and remarks.
KnupsENn, 1956, p. 525, plt. 2, fig. 7, off Cape Vert,
Senegal. Daurzenserc, 1889, p. 31, plt. 2, figs. 6a, 6b,
Azores.
Mitrolumna oliviformis CANTRAINE,; Locarp, 1892, p. 50,
“Mediterranean.”
Range: North Atlantic (Portugal, Nosre, 1932, to Sene-
gal, KNupsen, 1956) through the northern Mediterra-
nean (France, Bucguoy,DAUTZENBERG & Dot.Fus, 1883).
also reported as a fossil from’ Pliocene and Pleistocene
deposits of Europe. pa
Remarks: Mitrolumna has long been considered mono-
typic. Its type species, M. olivoidea, is now known to be the
last survivor of the genus with several species in the Old
World: Miocene and Pliocene. The discovery of a Recent
species in the Galapagos Islands referable to Mitrolumna
so far remote from the only other surviving ‘species seem-
ingly presents zoogeographic incongruities. Assuming the
correct generic assignment of the new Galapagan species,
one can speculate that these surviving species are relict
elements of a group that once enjoyed a much wider distri-
bution. It may eventually be found to have a geological
history similar to that of the related genus Mitromorpha
CarPENTER. Representatives of that genus are recorded
from the Pliocene and Pleistocene of California, Pliocene
of Florida, and Miocene of Florida and France (Powe LL,
1966), with 4 Recent species known from Californian and
west Mexican waters (KEEN, 1937; Coan, 1962) and at
least one species living in the western Atlantic (DALL,
1889).
The radular characters of the type species of Mitro-
lumna apparently have not been illustrated or described,
and we have not been successful in our efforts to extract
a radula from the very limited number of specimens that
were available for study.
Mitrolumna keenae EMERSON & RaDWIN, spec. nov.
(Plate 29, Figures 5, 6; Text figures 4, 5)
Diagnosis: Shell moderately large for the genus (attaining
18 mm in length), biconical; characterized by numerous
spiral incised lines on a buff-colored shell, with chestnut-
brown axial maculations between the spiral incised lines
which do not cross them.
Description: Shell moderately slender, whorls unshoul-
dered; spiral sculpture. of numerous fine, incised lines;
axial sculpture completely lacking; body whorl with 27-30
glyptae with interspaces varying in width. Immediately be-
low the suture there are 3 strongly incised lines alter-
nating with 3 grooves of typical strength forming a series
of 3 pairs of fine crowded striae; immediately below the
periphery of the body whorl is another series of 3 to 5
Page 154
THE VELIGER
Vol. 12; No. 2
crowded striae; remainder of the body whorl with regular
sculpture; spire consisting of 34 smooth, glassy, trans-
lucent-white nuclear whorls and 8 sculptured postnuclear
whorls which become progressively more strongly sculp-
tured posteriorly; outer lip thin with moderately devel-
oped anal notch, interior of outer lip with 14 short, weak
lirations. Exterior off-white with short chestnut-brown
maculations, darkest on the crowded subsutural inter-
spaces; a faint brown tinge on the peripheral group of
3 - 5 crowded interspaces; maculations randomly arranged
on the interspaces but do not transverse the spiral incised
lines.
The present species has a typical toxoglossate dentition
with a packet of 16 dart-like teeth, each about 10u long
(Text figures 4, 5) in a lateral pharyngeal pouch; the
packet remains cohesive during radular extraction as a
result of the presence of a series of filaments which con-
An 8
Mitrolumna keenae EMERSON & RADWIN, spec. nov.
Figures 4 and 5: Two views of radular tooth; greatly enlarged.
nect all the teeth together. A fragment of such a filament
may be seen in Text figure 4. This type of radular dentition
clearly shows the relationship of this species to others as-
signed to Mitromorpha, and other borsonine and mangel-
ine species (see Morrison, 1966; Powe 1, 1966).
Measurements: holotype 16.8 mm in length, 6.5mm in
greatest diameter; smallest paratype (from type locality)
13.3 mm in length; largest paratype 17.0 mm in length.
Type locality: near Tagus Cove, Isabella Island, Galapa-
gos Islands, dredged in 75-100 meters, January 25,
1968, by the DeRoys.
Type depositories: holotype, A.M.N.H. No. 152601
(Plate 29, Figure 5) ; 1 paratype, A. M. N. H. No. 152602
(Plate 29, Figure 6); 1 paratype, S.D.N.H.S. No. 50769;
1 paratype, A. D’Attilio Collection; 1 paratype, J. DeRoy
Collection. The paratypes are all from the type locality.
Remarks: All the specimens in the type lot were live-taken.
The holotype and one paratype (in the J. DeRoy Collec-
tion) are fully mature, whereas the paratypes in the A.
M.N.H., 8. D.N.H.S. and the A. D’Attilio Collections
are evidently immature.
The new species is generally similar to Mitrolumna
olwoidea but differs from it in the following ways: 1) its
shell is more than twice as large as that of M. olwoidea,
2) although both species have crowded spiral elements at
the periphery of the body whorl, the spiral elements on
spire whorls of M. olivoidea are strongly nodulose, where-
as those of MM. keenae have an almost planar surface,
3) the body whorl of M. olivoidea is broader than that
of M. keenae, imparting a stouter appearance to the shell
of the former, and 4) the two columellar plicae, typical
of the genus, are more strongly developed in M. olivordea.
Another genus that was given consideration for place-
ment of this new species is Artelia SHasxy (1961, p. 20,
pit. 4, figs. 7-9), type species A. mitriformis, by original
designation. Avielia, however, typically has a much nar-
rower shell, a more constricted body whorl, a much nar-
rower aperture, and spire whorls with less convex profiles
(Plate 29, Figure 1). Unfortunately, the radular charac-
ters of the type species of this eastern Pacific monotypic
genus are not known.
Explanation of Plate 29
Figures 1a, 1b: Arielia mitriformis SHAsKy, 1961, type species of
Arielia SHasky, 1961; 73-86 meters off Puerto Escondido, Baja
California, Mexico, “Puritan” Station 139; <3 (note predator’s drill
hole, Figure 1b).
Figures 3a, 3b:
Mitrolumna keenae EMERSON & RADWIN, spec. nov.
Figures 2a, 2b: juvenile specimen, paratype, with immature outer
lip and lacking pillar plaits; <3.
holotype, mature specimen; X3 (note artificial
“drill” hole, Figure 3b, drilled in order to remove soft parts for
radular study).
THE VELIGER, Vol. 12, No. 2 [EMERSON & Rapwin] Plate 29
Figure 1a Figure 1b
Figure 2a Figure 2b
Figure 3a Figure 3b
Vol. 12; No. 2
THE VELIGER
Page 155
ACKNOWLEDGMENTS
In addition to Mrs. Jacqueline DeRoy we are indebted to
the following friends and colleagues for courtesies of vari-
ous kinds: Mrs. Jean M. Cate, Dr. Eugene V. Coan, Mr.
Anthony D’Attilio, Mr. Harold S. Feinberg, Miss A. Ful-
lick, Dr. Maxime Glibert, Dr. Leo G. Hertlein, Miss Lynne
Judge, Dr. A. Myra Keen, Mr. William E. Old, Jr., Mr.
Richard E. Petit, Dr. Joseph Rosewater, and Dr. Donald
R. Shasky.
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Hinps, RICHARD BRINSLEY
1843. | On new species of Pleurotoma, Clavatula, and Mangelia.
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pls. 8-14 (Oct. 1844) ; pt. 3, pp. 49-72, pls. 15-21 (Jan. 1845)
[exact dates from a copy with original wrappers in the library
of the California Academy of Sciences].
Keen, A. Myra
1937. An abridged checklist and bibliography of west Ameri-
can marine Mollusca. Stanford Univ. Press, 87 pp.; 3 figs.
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi+624 pp.; 10
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1966. | West American mollusk types in the British Museum
(Natural History), II. Species described by R. B. Hinps.
The Veliger 8 (4): 265 - 275; plts. 46, 47; 6 text figs.
(1 April 1966)
KNUDSEN, JORGEN
1956. | Remarks on a collection of marine prosobranchs from
Senegal. Bull. Inst. Frang. Afr. Noire, Dakar, ser. A, Sci.
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1892. Les coquilles marines des cétes de France.
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Morrison, JosEPpH PauL ELDRED
1966. On the families of Turridae.
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1932. | Moluscos marinos de Portugal.
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1964. | Neogene mollusks from northwestern Ecuador.
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(October 1964)
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1860. | Nouveau supplément au catalogue des mollusques ma-
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Puitipr1, RUDOLF AMANDUS
1836. | Enumeratio Molluscorum Siciliae cum viventium tum in
tellure tertiaria fossilium quae in itinere suo observavit 1.
Berolini (Halis Saxonum). pp. i- xiv + 1 - 268; plts. 1-12
1844. Enumeratio Molluscorum Siciliae, cum viventium tum
in tellure tertiaria fossilium quae in itinere suo observavit 2.
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Pitspry, HENry Aucustus « AxEL ADOLF OLSSON
1941. A Pliocene fauna from western Ecuador. Proc. Acad.
Nat. Sci. Philadelphia 93: 1-79; plts. 1-19; 1 photograph
(9 September 1941)
PowELL, ARTHUR WILLIAM BADEN
1966. The molluscan families Speightiidae and Turridae.
Bull. Auckland Inst. & Mus., No. 5: 184 pp.; 23 plts.
REEVE, LovELL AUGUSTUS ;
1843[-1846]. Conchologia iconica: or illustrations of the shells
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THE VELIGER
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Rutscu, R.
1934. Die Gastropoden aus dem Neogene der Punta Gavilan
in Nord-Venezuela. Abhandl. Schweiz. Palaeont. Gesellsch.
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1836. | Catalogus conchyliorum Regni Neapolitani quae usque
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Suasky, DonaLp R.
1961. | New deep water mollusks from the Gulf of California.
The Veliger 4 (1): 18-21; plt. 4, figs. 1-10 (1 July 1961)
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1834. | Characters of new species of Mollusca and Conchifera,
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Sowerby, GeorcE BRETTINGHAM (2%? of name)
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Conch. 14 (8): 239 - 256
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1884.- Manual of Conchology. Philadelphia, ser. 1, vol. 6,
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Journ.
Vol. 12; No. 2
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Page 157
Escape Responses of Three Indian Molluscs
A. D. ANSELL
The Marine Station, Millport, Isle of Cumbrae, Scotland
INTRODUCTION
THE MARINE MOLLUSCs exhibit a wide range of defensive
responses to predators, including a variety of active mus-
cular reactions, often leading to flight by an accelerated
normal locomotory gait or by some mode of locomotion
not normally employed. In recent reviews of these respon-
ses FEDER & CHRISTENSEN (1966) and ANSELL (1969a)
have brought together a synopsis of many scattered refer-
ences to this phenomenon which clearly indicate that
specific responses are characteristic of certain families
or groups of families indicating a common and presumably
ancient origin for the response. Certain groups of the bi-
valves and gastropods thus apparently possess the poten-
tiality to respond to potential predators, and representa-
tives of these groups from different geographical areas or
habitats may also be expected to exhibit similar responses.
This is the case in the records presented here, which are
of interest since they relate to molluscs from the tropics,
while most of the previous records refer to species from
temperate seas. The responses were observed in the labor-
atory, in small aquaria, either at the Biological Oceano-
graphy Division of the Indian National Institute of
Oceanography, Ernakulam, South India, or at the Centre
for Advanced Studies in Marine Biology, Porto Novo.
OBSERVATIONS
Mactra olorina Putipri, 1846
(Bivalvia, Eulamellibranchia, Mactridae)
This species of Mactra, collected from sand near low
water from a beach near Shertallai, some 20 miles south
of Emakulam, exhibited the typical bivalve leaping re-
sponse when contacted on the siphon or mantle margins
by the foot of the predaceous gastropod Oliva gibbosa
(Born, 1778). The leaping response which occurred from
either the buried position or from the surface of the sand
was essentially the same as that described for the temper-
ate mactrid, Mactra corallina (LinNaEus, 1758) (AN-
SELL, 1969b). Repeated stimulation resulted after a short
time in a failure of the bivalve to respond by leaping,
although there was an increase in shell gape, a reaction
which together with the extension of the siphons to their
full length always occurs before leaping. The leaping
movements need not be described here since they show
no special features different from those of Mactra corallina.
Oliva gibbosa and Mactra olorina were both collected
from the beach at Shertallai, during January, 1968. The
general adaptations of these species to life on a surf-
washed tropical beach will be described elsewhere, but 1/7.
olorina is a fast burrowing active form whose normal
buried position is with the dorsal shell margins just covered
by the sand, while O. gibbosa is an active predator,
moving through or over the sand by means of a large
foot. At Shertallai, Oliva was found near high water mark
at low tide, and only during January of the period Janu-
ary to April, although it is not known whether its dis-
appearance later was caused by migration offshore, or by
local predation by the fishing community along the shore,
who utilize all types of shellfish food. Probably both play
a part, for CricHton (1942) records that on the east
coast of India near Madras, O. gibbosa is to be found
on the shore for a brief season only, early in February
each year. The feeding habits of the Olividae are not well
known. Members of the family are common in the faunas
of sand beaches in the tropics and warm temperate
zones, and it may be expected that they will be found to
elicit escape responses from molluscs and other inverte-
brates from such habitats. At Shertallai, other molluscs
collected from the sand included Donax incarnatus GMEL-
IN, 1791; the venerid Timoclea imbricata (J. DE C.
SoweErBy, 1826), Sunetta solanderu (Gray, 1821) and
the gastropod Bullia melanoides (DESHAYES, 1832). None
of these showed any response to O. gibbosa, but of these
Donax and Bullia are extremely active forms which exhibit
tidal migrations and hence any further specific response to
predators is probably unnecessary. The food of Oliva at
Shertallai could not be determined, but may include some
Donax which have been stranded by the receding tide,
since such stranded Donax and Oliva both occurred in a
narrow zone near the high water mark.
Page 158
Umbonium vestiarium (LinnaEus, 1758)
(Gastropoda, Prosobranchia, Trochidae)
The response of Umbonium, which was observed at Porto
Novo, was observed in an aquarium in which Umbonium
was present together with the carnivorous gastropod Bursa
Spinosa (LaMaRCK, 1816). These two species are not nor-
mally sympatric, and so these observations merely indicate
the ability of Umbonium to perform escape responses,
but do not indicate the predator which normally evokes
such responses. Umbonium responds by performing a series
of rolling movements produced by the action of the foot
which twists from side to side to push against the sub-
stratum after the animal has rolled over to lie on the
shell with the foot extended into the water. The move-
ments are essentially similar to those performed by some
members of the Trochidae, and especially Calliostoma
zizyphinum (LINNAEUS, 1758) (FEpDER, 1967; ANSELL,
1969a), in response to contact with certain asteroid star-
fish.
Umbonium is a common species on the surf-washed
sandy beaches near Porto Novo, and shows interesting
modifications of the basic trochid structure which enable
it to exploit this habitat. In the aquarium it exhibits
considerable activity, and movements of the same type as
the escape response occur apparently spontaneously. It is
likely that similar movements are used in active move-
ment on the shore, perhaps as in some bivalves represent-
ing a response to lack of suitable substrata for burrowing
(ANSELL, 1969b) or, as in the stenoglossan gastropod
Bullia melanoides, contributing to a complex behavioural
pattern by which the animal migrates up and down the
shore with each tidal cycle. In the aquarium similar
movements to those of the escape response also occur
when one Umbonium touches another while crawling and
similar intra-specific reactions were also noted for Bullia
melanoides from Shertallai Beach.
Babylonia spirata (LINNAEUS, 1758)
(Gastropoda, Prosobranchia, Buccinidae)
The conditions under which the response of Babylonia was
observed were also artificial and probably inno way repre-
sent a normal provocation. The specimen of Babylonia had
been placed for observation in a large glass vessel which
already contained a single individual of the volutacean
Harpa conoidalis Lamarck, 1822 and 2 Bursa spinosa.
After a short time the Bursa attacked the Harpa, which
retracted when pierced by the proboscis of the Bursa and
produced a copious secrétion of mucus. The Bursa was
disturbed and withdrew the proboscis, and during this
THE VELIGER
Vol. 12; No. 2
disturbance the response of the Babylonia, which was
nearby, was observed. The response consisted of the ani-
mal turning so that it lay on the side of the shell with the
foot extended in the water. A series of rolling /leaping
movements followed caused by a vigorous rotation of the
foot to press against the substrata first on one side and
then the other so that the shell is moved actively over the
surface in a zig-zag course. These movements are pro-
duced in the same way as, and are similar in essentials to,
the escape responses of the temperate buccinid Buccinum
undatum LINNAEUS, 1758 or of members of the family
Nassidae (FEDER, 1967). It is possible that some secretion
from the withdrawn proboscis was responsible for eliciting
the response.
DISCUSSION
These three examples of defensive responses by molluscs
from India present further evidence for the occurrence of
similar responses within certain groups of molluscs. Thus
the two gastropod groups represented, the Buccinacea and
the ‘Trochacea, together with the Zeugobranchia and
Patellacea, comprise a majority of the previous records of
locomotory responses by marine gastropods (ANSELL,
1969a). The members of the latter three groups are
almost exclusively inhabitants of rocky shores, and it is
therefore of considerable interest to find exactly similar
responses exhibited by a species which has become adapt-
ed to life in the disturbed environment of a tropical
surf-washed sand beach. Responses of this kind are appar-
ently of as much significance in the community relation-
ships of sand dwelling forms as they are in those of rocky
shores, which have provided most of the earlier examples.
This view receives support from the occurrence of a
response in the sandy-shore bivalve Mactra olorina. In
addition, this reaction is interesting since, although several
examples have been described of reactions by herbivorous
gastropods to the presence of, or contact with, carnivorous
gastropods, most previous records of leaping responses by
bivalves are of responses to asteroid starfish, the only ex-
ceptions of which I am aware being these of Ensts directus
(Conrad, 1843) which Turner (1955) described as
emerging from the sand in response to an attack by the
predaceous snail Polinices duplicata (Say, 1822) and the
reactions of Mactra sp. to Natica millepunctata LAMARCK,
1822 and N. hebraea and Spisula solida (LINNAEUS,
1758) to Lunatia nitida (Donovan, 1803) described by
Hirscw (1915) and ZiecELtmerer (1954) respectively.
There are no previous records of a member of the Oliv-
idae being responsible for eliciting escape responses in
other invertebrates.
Wal 1 2:No: 2
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Page 159
These three examples in part extend our knowledge of
defensive responses to a tropical fauna. The great variety
of species in tropical faunas and the resultant complexity
of the communities suggest strongly that such responses
may play an even more important role in community
interrelationships in the tropics than they do in temperate
waters. The numerical superiority of records of such re-
sponses from temperate forms probably reflects no more
than the greater opportunities at present available for
collection and observation of marine molluscs in temperate
regions.
ACKNOWLEDGMENTS
The observations were made during a collaborative pro-
gramme of study of sand beaches under the International
Biological Programme by the Marine Laboratory, Aber-
deen, and the Biological Oceanography Division of the
Indian National Institute of Oceanography, Ernakulam,
supported by a grant to Dr John Steele from the Royal
Society of London. I am grateful to Dr Steele for an
invitation to take part in this study and to the Royal Soci-
ety for its financial support. It is a pleasure to thank
Dr S. Z. Qasim, Director, Biological Oceanography Divi-
sion, Indian National Institute of Oceanography for
hospitality and for facilities provided at his laboratory,
and Professor R. V.Sheshaiya and Dr R. Natarajan of
the Institute of Advanced Studies in Marine Biology,
Porto Novo for their help in providing and identifying
an impressive collection of living molluscs during a very
short visit. I am also indebted to Mr H.E. J. Biggs,
British Museum (Natural History), who identified some
of the molluscs from Shertallai.
LITERATURE CITED
ANSELL, ALAN Davip
1969a. Defensive adaptations to predation in the mollusca.
Symp. Mar. Biol. Assoc. India (in press)
1969b. Leaping movements in the Bivalvia.
Soc. London (in press)
CricHton, MarsHALL DitwortTH
1942. Marine shells of Madras.
Soc. 42: 323 - 341
FeperR, Howarp MITCHELL
1967. | Organisms responsive to predatory seastars. Sarsia
29: 371 - 394
FepER, Howarp MitTcHELL & AAGE MoLLER CHRISTENSEN
1966. Aspects of asteroid biology. pp. 87-127 In: Physio-
logy of Echinodermata, RicHarp A. Boo.Lootian (ed.).
Intersci. Publ. (Wiley), New York.
Hirscu, Gottrwatt Cur.
1915. Die Emiahrungsbiologie fleischfressender Gastropoden.
Zool. Jahrb., Abt. Allg. Zool. 35: 357 - 504
Turner, Harry Jackson, Jr.
1955. How clam drills capture razor clams.
69 (1): 20-22
ZIEGELMEIER, ERICH
1954. | Beobachtungen tiber den Nahrungserwerb bei der Nati-
cide Lunatia nitida Donovan (Gastropoda Prosobranchia) .
Helgol. Wissensch. Meeresunters. 5: 1 - 33
Proc. Malacol.
Journ. Bombay Nat. Hist.
The Nautilus
Page 160 THE VELIGER Vol. 12; No. 2
Identification of the Feeding Types
in the Genus Conus LINNAEUS
BY
Cc. F LIM
Department of Zoology, University of Singapore, Singapore
(Plate 30; 5 Text figures; 2 Tables)
THE PREDATORY AND HIGHLY VENOMOUS gastropod genus
Conus has been established to be distinctly vermivorous,
molluscivorous, or piscivorous (KoHN, 1959a; ENDEAN & a2
Rupkin, 1965; Lim, 1968). It is also established that
these three feeding types in Conus can be ascertained by
examining the peculiarities in the anatomy of their radu-
lar teeth (ENDEAN & RuDKIN, op. cit.). So far no at-
tempts have been made to recognize the feeding types by
shell characters which would be convenient. It is the
intention here to try to do this, to try to assess the ratios
among the three feeding types and also to review the §
radular teeth characters in Conus.
THE RADULAR TEETH CHARACTERS
ENDEAN & RupkIN (1965) have already described the
diagnostic characters that could identify the feeding type
in the genus Conus. Basically, these are fairly reliable as
one might expect that the radular teeth would be well
adapted to the type of feeding. In review I am presenting
in Figure 1 an assortment of radular teeth to give an illus-
tration of the range of tooth-form existing in the genus.
Conus magus LINNAEUS, 1758, a widely distributed pisci-
vore in the Indo-Pacific region, possesses the typical form
of radular teeth for its feeding type (Figure 1, P 1). It is
(adjacent column —>)
Figure 1
Radular teeth in Conus
Pi: Conus magus LINNAEUS, 1758 (piscivore) P2: C. tulipa
LINNAEUS, 1758 (piscivore) Mu: C. textile LinNAEUS, 1758
(molluscivore) M 2: C. marmoreus LINNAEUS, 1758 (molluscivore)
V: C. betulinus LinnaEus, 1758 (vermivore)
b = barb; 1 = ligament; 's = shaft Scale = 1mm
Vol. 12; No. 2
a form similar to that seen in C. consors SowERBY, 1833,
C. monachus LINNAEUS, 1758, C’. purpurascens SOWERBY,
1833, and C. striatus LINNAEUS, 1758 (Lim, 1968; PEILE,
1939; Koun, 1959a). Conus tulipa LinNaAgEvs, 1758 (Fig-
ure 1, P 2), which is another piscivore, has radular teeth
which resemble the molluscivorous type more closely than
the typical piscivorous form. This is also true in the case
of C. geographus LINNAEUS, 1758, yet another piscivore
(ENDEAN & RuDKIN, op. cit.). The typical molluscivore
form of radular teeth is exemplified in C’.. textile LINNAE-
us, 1758, which is a widely distributed species in the Indo-
Pacific (Figure 1, M 1). A not-so-typical tooth but showing
all the molluscivore characters is seen in C’. marmoreus
Linnaeus, 1758 (Figure 1, M 2). Finally, in the vermi-
vores the radular teeth are rather uniform in structure as
in C. betulinus LinnNazus, 1758 (Figure 1, V). Teeth
similar to this have already been seen in C. caracteristicus
FiscHer, 1807, C. maldivus Hwass, 1792, C. monile
Hwass, 1792, C. striatellus Linx, 1807, C. coronatus
Gmeun, 1791, and many other vermivores (PEILE, 1939;
Koun, 1959a; and others).
RATIOS AMONG THE FEEDING TYPES
Fifty-five species of Conus from various parts of the world
were considered for the determination of ratios among
the vermivorous, molluscivorous, and piscivorous feeding
types (Table 2). They were chosen because of their feeding
types being known either from studies of their radular
teeth or gut contents or feeding observations. The results
of analyses revealed that 36 species were vermivorous, 9
molluscivorous, and 10 piscivorous. This works out to
65.46% vermivores, 16.36% molluscivores, and 18.18%
piscivores (Table 1). An approximation of this would be
a ratio of 4-5: 1: 1 for vermivores, molluscivores, and
piscivores respectively.
Table 1
Ratios of Conus species according to feeding-type
Number of Conus species considered 55
Percentage of vermivores (V) 65.46%
Percentage of molluscivores (M) 16.36%
Percentage of piscivores (P) 18.18%
Approximate ratios of V:M:P = 4-5:1:1
THE SHELL CHARACTERS
The most striking shell character of Conus consists of the
colour patterns or markings on the outer surface immedi-
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Page 161
ately beneath the periostracum. There is a great variety
of these markings, and attempts to sort the feeding types
of the species proved difficult. As a result of extensive
studies of the shell characters of species of known feeding
types (by examinations of their radular teeth, gut contents,
or feeding observations) I have chosen 3 characters that
can be used to some degree for diagnosing feeding type.
They are pink tip (PT) of the spire, tent marks (TM)
on the body whorl (Plate 30, Figures A to D), and inter-
rupted striae (IS) on the body whorl (Plate 30, Figures E
to H). Analyses of these 3 characters in 55 species are
shown in Table 2. It can be observed that none of the
characters is exclusive to any of the feeding types. But
they do show that each of the 3 characters is preponder-
ant to one feeding type only. Graphic presentations illust-
rated clearly that PT occurred in 100% of the piscivores,
44.4% in the molluscivores, and 5.6% in the vermivores
considered (Figure 2). The TM character was present in
Vv
Figure 2
Graphic presentation of pink tip (PT) character in Conus shaded
black. Vermivores (V) 5.6%; molluscivores (M) 44.4%;
piscivores (P) 100%
100% of the molluscivores, 10% of the piscivores, and
2.7% of the vermivores considered (Figure 3). Thirdly,
the IS character was positive in 100% of the piscivores,
13.9% of the vermivores, and totally absent in the mol-
luscivores considered (Figure 4). From these analyses it
may be concluded that TM is positively relevant to mol-
luscivores and PT and IS are to piscivores; and that TM,
PT, and IS are mostly negative to the vermivores.
If these conclusions are to serve any useful purpose,
these characters are examined closely in Table 2. It can
be observed that all the piscivores considered have the
combination of PT and IS characters, except Conus geo-
Page 162
Figure 3
Graphic presentation of tent marks (TM) character in Conus
shaded black. Vermivores (V) 2.7%; molluscivores (M) 100%;
piscivores (P) 10%
graphus, which has the characters of PT and TM. Conus
geographus can easily be distinguished from all other spe-
cies of Conus by its possession of a wide shell aperture.
The transverse width of the aperture at the waist is more
Vv
Figure 4
Graphic presentation of interrupted striae (IS) character in Conus
shaded black. Vermivores (V) 13.9%; molluscivores (M) 0%;
piscivores (P) 100%
THE VELIGER
Vol. 12; No. 2
than twice that of the ventral body whorl at the same
level (see Figure 5). Summarizing, the piscivores can be
differentiated from either the vermivores or molluscivores
by the following key.
Figure 5
Measurement of transverse width of shell aperture (b) at the waist
(x), and width of ventral body whorl (a) at same level
KEY to PISCIVOROUS Conus
(to be attempted on shells that are completely clean of
periostracum )
1. Transverse width of shell aperture at waist (see Figure
5)more than twice the transverse width of ventral
body whorl at the same level. Shell spire with pink
tip". Ae ee ee piscivore (part)
— ‘Transverse width of shell aperture less than twice that
of ventral body whorl at the same level. Shell spire
with) or without pink tip) ieee cena eee 2
2. Body whorl of shell with transverse interrupted striae
or striations on almost all its length; without tent
MALKS hes ccstsntsecnaabeennren acacia ae 3)
— Body whorl of shell without such markings; with tent
marks molluscivore or vermivore (part)
3. Shell spire with pink tip piscivore (part)
— Shell spire without pink tip vermivore (part)
Explanation of Plate 30
The tent marks (TM) and interrupted striae (IS) characters on
the body whorl of the shell in Conus.
A: Conus amadis GMELIN, 1791; B: C. marmoreus LINNAEUS, 1758;
C: C. aulicus LINNAEUS, 1758;
(A to D: molluscivores)
D: C. textile LinNaEus, 1758;
E: C. tulipa LinnaEus, 1758; F: C. stercusmuscarum LINNAEUS,
1758; G: C. striatus LINNAEUS, 1758; H: C. magus LINNAEUS, 1758;
(E to H: piscivores)
[Lim] Plate 30
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Vol. 12; No.
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Page 163
Table 2
Shell markings of Conus species of known feeding type
Reference to radular teeth/gut
Species of Conus PT T™/ IS V M P contents/feeding observations
1. amadis GMELIN, 1791 — + — — + — Penne, 1939
2. ammuralis LINNAEUS, 1758 + + — — + — ENDeEaAN & Rupkxin, 1965
3. arenatus Hwass, 1792 — — + + — — ENDEAN & Rupkin, 1965
4. aulicus LINNAEus, 1758 — + — — + — EnpeEaN & Rupkin, 1965
5. betulinus LINNAEUS, 1758 —- — — + — —
6. californicus Hinps, 1844 — — — + — — WHYSNER & SAUNDERS, 1963
7. caracteristicus Fiscuer, 1807 — +-—- + — —
8. catus Hwass, 1792 + — + — — +. Koun, 1956
9. chaldeus Ropinc, 1798 — — — + — — Koun, 1959a
10. consors SowERBy, 1833 + — + — — + Lin, 1968
11. coronatus GMELIN, 1791 — — + + — — Koun, 1959b
12. dalli StEaRNS, 1873 — + — — + — Nysaxken, 1968
13. distans Hwass, 1792 — — — + — — Kosn, 1959a
14. ebraeus Linnaeus, 1758 — — — + — — Konn, 1959a
15. eburneus Hwass, 1792 — — — + — — Konn, 1959a
16. emaciatus REEVE, 1849 + ENDEAN & RupkiIn, 1965
17. figulinus LinnaEus, 1758 SS ae) KOHN, 1959)b
18. flavidus LAMarck, 1810 — — — + — — Koun, 1959a
19. geographus LinNaEus, 1758 + + — — — + Enpean « Rupxin, 1965
20. glans Hwass, 1792 + — — + — — Koun, 1959b
21. wnperialis LINNAEus, 1758 — — + + — — Konan, 1959a
22. leopardus Ropine, 1798 — + Koun, 1959 a
23. litteratus LinNAEus, 1758 — — — + — — EnpeAN « Rupkin, 1965
24. lividus Hwass, 1792 + — — + — — Koun, 1959b
25. luteus SowErsy, 1833 — — + + — —
26. magus Linnaeus, 1758 + — + — — + Enpean « Rupxin, 1965
27. maldivus Hwass, 1792 — — — + — — Pete, 1939
28. marmoreus LINNAEUS, 1758 — + — — + — Koun, 1959a
29. mediterraneus Hwass, 1792 — — — + — — Arprrs, 1932
30. miles LINNAEUS, 1758 —_—- — + — — Koun, 1959a
31. monachus LinNAEus, 1758 + — — — +
32. monile Hwass, 1792 — — — + — — Pete, 1939
33. moreleti Crosse, 1858 — — — + — — Koun, 1959a
34. musicus Hwass, 1792 — — + + — — Konn, 1959 a (as C. abbreviatus REEVE)
35. obscurus SowERBY, 1833 + — + — — + Koun, 1959a
36. omaria Hwass, 1792 + + — — + — _ EnNDEAN & Rupkin, 1965
37. pennaceus Born, 1778 + + — — + — Koun, 1959a
38. pertusus Hwass, 1792 — — — + — — Koun, 1959a
39. planorbis Born, 1780 — — — + — — ENpDEAN & Rupkin, 1965
40. pulicarius Hwass, 1792 — — — + — — Koun, 1959a
41. purpurascens SowERBy, 1833 > — + — — + Petre, 1939
42. quercinus LicHTFooT, 1786 SS Koei, IED a
43. rattus Hwass, 1792 — — — + — — Koun, 1959a
44. retifer MENKE, 1829 > + — — + — Koun, 1959a
45. sponsalis Hwass, 1792 — -— —— —«s*t — — Konwn, 19594
46. stercusmuscarum LinnaEus, 1758 + — + — — +. EnNpeaN & Rupkin, 1965
47. striatellus Linx, 1807 — — — + — — Prez, 1939
48. striatus LINNAEUS, 1758 + — + — — + Koun, 1959a
49. terebra Born, 1780 — — — + — — Koun, 1959b (as C. clavus LinNaEvus)
50. tessulatus Born, 1778 p=
51. textile LINNAEUuS, 1758 — + — — + — Konn, 1959b
52. tulipa LinNAEUus, 1758 + — + — — + Konn, 1963
53. vexillum GMELIN, 1791 — — — + — — Konan, 1959a
54. virgo LinnaEus, 1758 — — — + — — Konan, 1959b
55. vitulinus Hwass, 1792 — — — + — — Konun, 1959a
IS = interrupted striae; M = molluscivore; P = piscivore;
PT = pink tip; TM = tent marks; V = vermivore
Page 164
THE VELIGER
Vol. 12; No. 2
DISCUSSION
The radular tooth characters previously described by
ENDEAN & RupkKIN (1965) may be sufficiently diagnostic
in determinations of feeding type. But this may not al-
ways be convenient. It is considered that shell characters,
if workable, would be preferable. Although the shell
characters presented here cannot be used for determin-
ing all 3 feeding types of Conus species, it may be employed
to distinguish the piscivores from either the molluscivores
or vermivores. It has already been known that the pisci-
vores are the only species that may cause any serious
danger to human beings (ENDEAN & RUDKIN, of. cit.).
In this respect the proposed diagnosis by shell characters
may be of interest.
The analysis for the ratios among the 3 feeding types
indicates clearly that the vermivores (65.46%) far out-
number either the molluscivores (16.36%) or piscivores
(18.18%). Studies of 50 species of Conus from Singapore
and the Malay Peninsula revealed 38 (76%) to be vermi-
vores, 7 (14%) molluscivores, and 5 (10%) piscivores
(Lim «& Tan, in press). Comparing the ratios of feeding
type in species from one area and that of feeding types in
species from numerous areas combined show no highly
marked differences.
ACKNOWLEDGMENTS
May I add an expression of thanks to Mr. A. J. Johnson
and Mr. J. R. Fisher for kindly allowing me to examine
their joint collection of cone shells.
LITERATURE CITED
ALPERS, FE
1932. Uber die Nahrungsaufnahme von Conus mediterraneus
Brug., eines toxoglossen Prosobranchiers. Publ. Staz. Zool.
Naples 11: 426 - 445
ENDEAN, Ropert & CLARE RUDKIN
1965. Further studies on the venoms of Conidae.
2: 225 - 249
Koun, ALAN JAcoss
1956. Piscivorous gastropods of the genus Conus. Proc.
Natl. Acad. Sci. U.S.A. 42: 168 - 171
Toxicon
1959a. The ecology of Conus in Hawaii. Ecol. Monogr.
29: 47 - 90
1959b. Ecological notes on Conus (Mollusca:Gastropoda) in
the Trincomalee region of Ceylon.
(13) 2: 309 - 320
1963. | Venomous marine snails of the genus Conus. In:
H. L. Kegcan & W. V. MacrarLane, ed., Venomous and poison-
ous animals and noxious plants of the Pacific region. Pergamon
Press, London, pp. 83 - 96
Lim, C. FE
1968. A comparative analysis of the feeding adaptations in the
predatory gastropod Conus LINNAEUS. Third Europ. Mala-
col. Congr. Vienna (abstract)
Lim, C. EF « W.C. Tan
(in press) Size and metabolic rate in the venomous snail Conus
LINNAEUS.
NyYBAKKEN, JAMES WILLARD
1968. Notes on the food of Conus dalli Stearns, 1873.
The Veliger 11 (1) : 50 (1 July 1968)
Ann. Mag. Nat. Hist.
Pee, A. J.
1939. Radula notes, 8 Conus. Proc. Malacol. Soc. London
23: 348 - 355
Wuysner, J. A. & Paut R. SAUNDERS
1963. Studies on the venom of the marine snail Conus cali-
Toxicon 1: 113 - 122
fornicus.
Vol. 12; No. 2 THE VELIGER Page 165
Panamic Sites and Archaeological Mollusks of Lower California
BY
LAWRENCE H. FELDMAN
Department of Anthropology, Pennsylvania State University, University Park, Pennsylvania 16802
(2 Tables)
INTRODUCTION
THIS PAPER IS AN ATTEMPT to systematically list all Lower
California archaeological molluscan species from assemb-
lages that are primarily Panamic in origin. Therefore the
series of middens on the Pacific coast of the peninsula,
north of Punta San Antonio, containing Californian faun-
al province shells (c. f Moriarty, 1968a), are not men-
tioned in this paper. Table 1 indicates what species are
found at each site and Table 2 tries to show what the
species tell us about the site occupations.
On the basis of Table 2 we might divide the Panamic
sites into two groups. A northern group is represented by
the sites of Playa de Moreno, San Felipe and Bahia de los
Angeles. These were inhabited only in the wet winter
months; as for example at San Felipe, where the Kiliwa
in historic times went to fish. “By summer, it was said, the
fresh water at San Felipe was so dried up that no one
lived there permanently. The indians gathered clams and
mussels and caught fish and dried them in the sun to
take back to their permanent rancherias,” (Meics, 1939:
p. 27). It was only at sites of this part of Lower Califor-
Table 1
Archaeological Mollusca
PELECYPODA
Pecten vogdesi ARNOLD, 1906
Dosinia ponderosa (Gray, 1838)
Chione undatella (Sowersy, 1835)
Pinna rugosa SowERBy, 1835
Cardita affinis Sowersy, 1833
Ostrea palmula CarPENTER, 1856
Anomia peruviana b’OrzIGNy, 1846
Anadara multicostata (SoweErRBy, 1833)
Ostrea fisheri DAL, 1914
Pteria sterna (Goutp, 1851)
Trachycardium panamense Dau, 1916
Laevicardium elatum (Sowersy, 1833)
Chione gnidia (BropERIP & SoweErRBY, 1829)
Chione fluctifraga (SowerBy, 1853)
Spondylus princeps Broverip, 1833
Protothaca grata (Say, 1831)
Tagelus californianus (Conran, 1837)
Ostrea angelica RocHEBRUNE, 1895
Glycymeris gigantea (Reeve, 1843)
Megapitaria squalida (Sowersy, 1835)
SO He SOS CCAS sie Sik
3 il, 2, B C
1
x ls 2 il, 2 R
SK 1 52
1 R
A 1
1 A
1 1
52 1
1 A x 3
x 1 2
x 1 2
2 A
il, 2 iP 2,
1 2
1 ?2 A
A 33, 2 R
P 2 C
Page 166
THE VELIGER
Vol. 12; No. 2
Table 1 [continued]
Chama buddiana C. B. Apams, 1852
Chione cortezi (CARPENTER, 1864)
Mulinia pallida (BropERip & SowERBY, 1829)
Aequipecten tumbezensis (p’OrBIGNY, 1846)
Mulinia coloradoensis Datu, 1894
Ostrea iridescens Hanuey, 1854
Trachycardium senticosum (SOWERBY, 1833)
Semele flavescens (Goutp, 1851)
Pinctada mazatlanica (HaNLEy, 1856)
Arca pacifica (SowERBY, 1833)
Barbatia reeveana (p’Orxicny, 1846)
Anadara formosa (SowErRBY, 1833)
Glycymeris maculata (BroperipP, 1832)
Glycymeris multicostata (SOWERBY, 1833)
Modiolus capax (Conrap, 1837)
Aequipecten circularis (SowERBY, 1835)
Lyropecten subnodosus (SoweErRBy, 1835)
Trigoniocardia biangulata (BRoDERIP & SOWERBY, 1829)
Periglypta multicosta (SowERBY, 1835) —
GASTROPODA
Haliotis sp.
Bulla gouldiana Pirssry, 1895
Fusinus dupetitthouarsi (K1ENER, 1846)
Hexaplex erythrostomus (Swainson, 1831)
Crucibulum spinosum (SoweErsy, 1824)
Conus regularis SowERBY, 1833
Strombus gracilior SowERBy, 1825
Turbo fluctuosus Woop, 1828
Tegula rugosa (A. ApaMs, 1855)
Olivella biplicata (SowERBy, 1825)
Polinices reclusianus (DESHAYES, 1839)
Muricanthus nigritus (PHipri, 1845)
Strombus galeatus Swainson, 1823
Conus fergusoni SowERBy, 1873
Oliva incrassata (LicHTFooT, 1786)
Cerithium stercusmuscarum VALENCIENNES, 1833
Crepidula incurva (Broverip, 1834)
Calyptraca mamillaris BRoDERIP, 1834
Crucibulum umbrella (DEsHaAyYES, 1830)
Calyptraca subreflexa (CARPENTER, 1856)
Nerita funiculata MENkE, 1851
Acmaea mitella MENKE, 1847
PM SF? SLG@ GC? BAS (S)iceesie
73)
x
x
x
4
2,
2
2
5
1 C
Cc
R
1 R
Cc
1 P R
A
P A
R
R
3
1
1
52
x ey 1 R
1
A 12
1
1 1
3
f 1 iP 2
1 A 2
Pp R 2
1
1 4
2 R
x
x
x
2
2
ee
1 Playa de Moreno Beach, north of San Felipe. Collected by
James R. Moriarty on October 31, 1958. Probably mixed with
modern shell.
2 San Felipe. (1): ScHENck & GirForp, 1952. (2): Collected by
James R. Moriarty from beach on October 31, 1958. Sample
probably mixed with modem shell. (3): Clam shell from ex-
tensive superficial midden on beach ridge, at edge of coastal
terrace, just above aud W of air strip occupying a stranded
lagoon, close to the San Felipe village well (31°01’N; 114°51’
W), Moriarty, 1968a.
3 San Luis Gonzaga Bay, Coan, 1965. (P) = present; (C) =
common; (A) = abundant.
4 Gardner Cave, MeicHan, 1965.
5 Bahia de los Angeles. (1): E.L. Davis correspondence (July
Vol. 12; No. 2 THE VELIGER Page 167
1965). (2): midden material collected by James R. Moriarty.
(3): cave and midden material, Massey « Osporne, 1961.
® San José Island, Emerson, 1960. (R) = rare; (C) = com-
mon; (A) = abundant.
7 San Lucas. (1): a 30mm diameter ring with a 10mm hole that
was ground down and polished; collected by H.N. Lowe at
Cape San Lucas and at present located in the San Diego Natur-
al History Museum collections. (2): shells from terrace along
nia that trade shells from other faunal provinces appear
(1. e. Haliotis sp., Olivella biplicata).
The San José Island site represents a possible southern
group of summer occupation sites. Like the San Blas
Nayarit shell middens, free swimming pectens are very
Golfo de California at Punta Pescadero (23°47’06” N; 109°42’
00” W), Mortarty, 1968. (3): Shell from shore of Ensenada
de los Aripes, at Radio Station of La Paz, W of town proper,
in indurated sand below high-tide level, and below level of
adjacent midden (ca. 24°09’ N; ca. 110°19’30” W), Morrarty,
1968. (4): Punta Conejo, a shell mound about 40 miles N of
La Paz and } to 4 mile inland from Punta Conejo on road to
La Paz; collected by C. W. Meighan. (5): Cape San Lucas
area shells, MAssEY « Osporne, 1961.
abundant and suggest the presence of a deep water fishing
industry (FELDMAN, n.d.). Perhaps a summer time sea
salt production was also a factor in the occupation of the
San José Island and San Blas sites at this season (c. f
Moriarty, 1968b).
Table 2
Archaeological Mollusca Sites
Site® Radiocarbon Date® Species Niche’? —_ Seasonality"
Playa de Moreno — ResiSc35 Bell ?winter only
Podrs laisse ||
San Felipe (1) — IRe7/ 2 SoilOe Teil s winter only
?:1;R,S8:4
San Felipe (2) — RESIS es eos —
R, S:1
San Felipe (3) 598 A. D. R:1 =
San Luis Gonzaga = Rell e Seite winter &
R, 8:2 summer
Gardner Cave 1352 A.D. SeBo Peil —
Bahia de los Angeles (1) — Reo SEOs Bele winter only
PHO Sells lasyells
S, B:2
Bahia de los Angeles (2) 4138 B. C. RESTS. ope Bel winter only
532 B.C. Pail Si asyell Bish}
Bahia de los Angeles (3) — S45 FSslle Psi
San José Island = Re Po Ge Teil ¢ summer only
?:6;R,S:4; FS:2
San Lucas (1) — R:1 —
San Lucas (2) 263 A. D. R:1 8:2 —
San Lucas (3) 1152 B.C. S:l a
San Lucas (4) — S:1 —
San Lucas (5) — eal ase
8 See footnotes of Table 1 for an explanation of site numbers. At B
9 Except for the Gardner Cave date, all Radiocarbon dates are FS = free swimming animal;
? = niche not known;
from Moriarty, 1968.
'o Based upon data in Parker, 1964, and Kren, 1958:
B = low-salinity lagoon and mangrove environment (brack-
ish) ;
S = intertidal sand beaches and sand flats to 26 meters 5
R = intertidal and shallow rocky shore assemblage;
the numbers denote how many species are of each niche per
site assemblage.
Seasonality data, from Marxrin, 1962, are based upon recent
collecting habits of the Seri Indians of Sonora for Gulf of
California species.
Page 168
THE VELIGER
Vol. 12; No. 2
Panamic Lower California shows continuous exploita-
tion of its molluscan resources from 6000 years ago up to
the arrival of the European explorers. However, still little
is known about the archaeological mollusks of most of
the peninsula. Regions like the coast of the Bahia de
Sebastian Vizcaino, where the Panamic fauna presumably
grades into the Californian fauna and the peninsula
from La Paz to Cabo San Lucas are almost totally
unknown. Species lists from the middens of these and
other Lower Californian areas might be of considerable
interest for the light they could shed on the way of life of
the aboriginal inhabitants of the peninsula.
ACKNOWLEDGMENTS
I would like to express my thanks to E. P. Chace, C. W.
Meighan, J. R. Moriarty, and E. L. Davis for making
available unpublished data on the archaeological shells
of Lower California.
LITERATURE CITED
Coan, EucGENE VICTOR
1965. Kitchen midden mollusks of San Luis Gonzaga Bay.
The Veliger 7 (4) : 216 - 218; plt. 28; 1 table (1 April 1965)
EMERSON, WILLIAM KEITH
1960. Results of the Puritan-American Museum of Natural
History Expedition to Western Mexico, 12: Shell middens of
San José Island. Am. Mus. Novitates, No. 2013: 1-9,
4 figs. (18 August 1960)
FELDMAN, LAWRENCE H.
no date. | Greater Nayarit archaeological mollusks and archaeo-
molluscan units in northern Mesoamerica. Paper prepared
for inclusion in C. W. MEIcHAN’s monograph on the site of
Amapa, Nayarit (12 December 1968)
Keen, A. Myra
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi + 624 pp.; illus.
Stanford, Calif. (Stanford Univ. Press)
Mackin, B.
1962. Seri Ethnozoology.
7: 1-56
Massey, WiiuiaM C. & CaRroLyn M. OssBorNE
1961. A burial cave in Baja California. The Palmer Collection,
1887. Univ. Calif. Anthrop. Records 16 (8) : 338 - 353
MEIGHAN, CLEMENT
1965. Cave paintings of Baja.
Occ. Pap. Idaho State Coll. Mus.
Desert 28 (7): 16-19
(July 1965)
Meics, PEVERIL
1939. The Kiliwa Indians of Lower California.
Americana 15. Univ. Calif. Press, Berkeley, Calif:
Moriarty, JAMES ROBERT
1968a. Climatologic, ecologic and temporal inferences from ra-
diocarbon dates on archaeological sites, Baja California, Mexico.
Pacif. Coast Archaeol. Soc. Quart. 4 (1): 11-38
(January 1968)
1968b. The socio-political and economic influences related to the
production and distribution of salt. Anthrop. Journ. Can.
6 (1): 2-15
Parker, Rosert H.
1964. | Zoogeography and ecology of some macro-invertebrates,
particularly molluscs, in the Gulf of California and the conti-
nental slope off Mexico. Vidensk. Medd. Dansk naturh.
Foren. 126: 1 - 178; 15 plts.; 29 text figs.; 7 tables (17 Febr. ’64)
ScuHEnck, W. EcBerT « E. W. Girrorp
1952... Archaeological sites on opposite shores of the Gulf of
California. Am. Antiquity 17 (3): 265. (January 1952)
Ibero-
Vol. 12; No. 2
THE VELIGER
Page 169
Cyclostrema miranda BARTSCH,
a Synonym of Jornus subcarinatus MONTAGU
DONALD R. MOORE
Institute of Marine Sciences, University of Miami, Miami, Florida 33149
(3 Text figures)
Paut BartscH, during the early years of the present
century, described a large number of micromollusks from
the Pacific coast of the United States. A number of these
species were assigned to the Vitrinellidae although a mod-
ern classification would put them in two orders and several
families. One of the species described by Bartscu (1910)
was Cyclostrema miranda, a small depressed gastropod
about 2 mm in diameter. Since its description, it has not
been reported from elsewhere on the Pacific coast since
Datu (1921) and Keen (1937) both list it from only the
type locality, San Pedro, California.
In December, 1965, I examined the type specimens of
west coast Vitrinellidae described by Bartsch. His Cyclos-
trema miranda was of special interest since his illustration
showed a shell with a very large oblique aperture. Typical
Vitrinellidae do not have such a widely open aperture,
although Macromphalina, a genus that does, has been
placed in the family by some authors.
When I examined the holotype of Cyclostrema miranda,
I saw that it was a specimen of Tornus subcarinatus
(Monrtacu, 1803). To make certain, I compared it with
specimens in the JEFFREYS collection from Guernsey, one
of the Channel Islands between England and France.
This species is the type of the genus Jornus, and hence of
the family Tornidae. It is thus of more than usual interest
to those who work on minute mollusks. Tornus subcarina-
tus is a European species ranging from Ireland and south-
ern England (Forses & HANLEy, 1849), southward in the
Atlantic to Senegal and Gabon (Nick és, 1950), and it
is also found in the Mediterranean at least as far east as
the Adriatic (Brusina, 1866).
Now that the identity of the species is established, it
would be of interest to know if it is actually living on the
* Contribution No. 1085 from the Institute of Marine Sciences,
University of Miami, Miami, Florida 33149
California coast. BartscH (1910) states that the type,
and two other specimens in Mrs. Oldroyd’s collection,
were collected by Mrs. Oldroyd at San Pedro, California.
Ordinarily, this locality might be accepted at face value,
but the lack of additional material, in a well collected
area, is enough to instill doubt. Another incident that
4
Figures 1 to 3
Top, bottom, and lateral views of Tornus subcarinatus
(after BARTSCH, 1910)
may have some bearing on the case was the discovery of
two specimens of Tornus subcarinatus in the Stimpson
Collection at the University of Miami. The locality on the
label was “Dry Tortugas,” a small group of islands near
Key West, Florida. Charles T. Simpson was a scientific
collector with a special interest in the Mollusca of Florida,
and was a contemporary of Mrs. Oldroyd. It is possible
that someone, through either duplicity or ignorance, gave
or sold specimens to both collectors with false locality
Page 170
THE VELIGER
Vol. 12; Now 2
data. Tornus subcarinatus is a shallow water animal, and
it is highly unlikely that it could be living on both sides of
North America without being reported from more than a
single locality. I personally have examined many shallow
water bottom samples from the Florida Keys and adjoining
areas without finding the species.
The family Tornidae is equated with the Vitrinellidae
by some workers, while others consider them to be distinct.
I discussed this situation in 1965. My views have been
modified slightly since some vitrinellids have a very long
gill which protrudes a little beyond the edge of the aper-
ture. However, it appears to be somewhat different from
the gill of Tornus subcarinatus as figured by WoopwarpD
(1899). The presence of a penis in vitrinellidae is based
on observations by FRETTER (1956) on Czrculus striatus
(Puiuippr) and by Moore (1964) on Parviturboides inter-
ruptus (C.B. ApAMs). (Woopwarp (1899) did not find
a penis in the male T: swbcarinatus that he examined. We
still lack comparative data, but it does not appear that
Tornus and the vitrinellids should be placed in the same
family. One fundamental difference is the operculum, for
even very depressed vitrinellids with a very oblique aper-
ture, such as Cochliolepis parasitica Stimpson, have a
circular multispiral operculum. Tornus subcarinatus, on
the other hand, has an oval paucispiral operculum.
Apparently, then, the Oldroyd and Simpson specimens
of Tornus subcarinatus have spurious locality data and
probably originally came from somewhere in the eastern
hemisphere. There are apparently no congeneric species
in the western hemisphere, but, until we know the fauna
better, the presence of some member of the family cannot
be ruled out.
ACKNOWLEDGMENTS
I am grateful to the Division of Mollusks, U.S. National
Museum, for the opportunity to examine the type speci-
mens of species described by Paul Bartsch. This work was
supported by National Science Foundation Grants GB-
3104 (UM 8190) and GB-5055 (UM 8753).
LITERATURE CITED
BartscH, PauL
1910. Descriptions of new mollusks of the family Vitrinellidae
from the west coast of America. Proc. U.S. Nat. Mus.
39 (1785) : 229 - 234; plts. 39, 40
Brusina, S.
1866. | Contribuzione della fauna dei molluschi dalmati.
Soc. I. R. Zool. Bot. 17: 1- 134; 1 plt.
Dati, WILLIAM HEALEY
1921. Summary of the marine shellbearing mollusks of the
northwest coast of America, from San Diego, California, to
the Polar Sea, mostly contained in the collection of the
United States National Museum, Smithson. Inst., U.S.
Nat. Mus. Bull. 112: 1-217; plts. 1-22 (24 February 1921)
Forses, EpwarD & SYLVANUS HANLEY
1853. A history of British Mollusca, and their shells. Van
Voorst, London. 2: 1-557
FRETTER, VERA
1956. The anatomy of the prosobranch Circulus striatus (PHIL-
IpPi) and a review of its systematic position. Proc. Zool.
Soc. London 126 (3) : 369 - 381; figs. 1-5
Keen, A. Myra
1937. An abridged checklist and bibliography of west North
American marine mollusca. Stanford Univ. Press, Stanford,
Calif., pp. 1 - 87
Moore, Dona.p R.
1964. The family Vitrinellidae in South Florida and the Gulf
of Mexico. Ph. D. dissertation, Univ. Miami: 1 - 235;
figs. 1-35
1965. New species of Vitrinellidae from Gulf of Mexico and
adjacent waters. The Nautilus 78 (3): 73-79; 9 figs.
Nick.es, M.
1950. | Mollusques testacés marins de la céte occidentale d’Af
rique. Man. ouest-afric. 2: 1-269; figs. 1-459. Paris,
Lechevalier.
Woopwarpb, Martin FOUNTAIN
1899. On the anatomy of Adeorbis subcarinatus Montacu.
Proc. Malacol. Soc. London 3: 140 - 146; plt. 8
Vol. 12; No. 2
THE VELIGER
Page 171
Relative Growth Patterns of Two West Coast Squid
(Gonatus fabric and Gonatopsis borealis)
BY
LARRY T. SPENCER
Plymouth State College,
Plymouth, New Hampshire 03264
(2 Text figures)
INTRODUCTION
THE OPTIMAL METHOD of determining patterns of growth
in any organism is to maintain the species in a laboratory
situation where measurements of size increases can be
made throughout the lifetime of the individuals. For many
organisms this task is quite impossible. Therefore measure-
ments of relative growth are quite often used as substitutes.
Although relative growth patterns do not indicate the
time taken to attain maximum size, they are useful to
compare the growth of two different species of squid.
HarFner (1964)utilized the differences in growth pat-
terns to taxonomically differentiate between Loligo pealei
and Lolliguncula brevis in Chesapeake Bay, Maryland.
This study presents the differences in relative growth of
Gonatus fabric (LICHTENSTEIN, 1818) and Gonatopsis
borealis Sasaxt, 1923, two pelagic squid species of the open
ocean off Oregon.
METHODS
Eighty-one Gonatus fabricii and 88 Gonatopsis borealis
were examined in detail. The specimens measured includ-
ed those collected by the Department of Oceanography,
Oregon State University and others loaned to the depart-
ment by the U.S. Fish and Wildlife Service and the Uni-
versity of Washington (Pearcy, 1965). Eight morpholog-
ical characteristics were measured (these are described
by Harrner, 1964): mantle width (MW); head width
(HW); head length (HL); nuchal cartilage length (NCL);
fin width (FW); fin length (FL); siphon width (SW) ;
and siphon length (SL).
' This work was done as a Graduate Student in the Department of
Zoology, Oregon State University, Corvallis, Oregon 97331
The relative growth patterns were determined as fol-
lows:
1. The squid were grouped in 10mm intervals of dorsal
mantle length. The average measurement for each of
the 8 characters was calculated and in turn divided by
the average dorsal mantle length for that interval. The
resultant figure is defined as the morphometric index.
2. The indices for each size interval were plotted against
average dorsal mantle length and the points were con-
nected. The resultant curve is defined as the growth
pattern.
The curves for Gonatus fabricitt are more accurate in
the lower size range (15 - 60mm) than in the higher size
range (60 - 190mm) because of a lack of larger individ-
uals. Comparable numbers of Gonatopsis borealis were
measured in each size interval.
RESULTS
Gonatopsis borealis
The relative growth patterns of Gonatopsis borealis are
presented in Figure 1 (left). Two distinct patterns are
readily discernible. One pattern is a rapid increase of the
character size up to 30mm mantle size followed by either
a plateauing (FL) or a continued peaking and then
decrease (FW). The other pattern is a relatively rapid
decrease of the character changing at 30mm mantle
length to a less rapid decrease (MW, HW, HL, SW, SL,
and NCL). Of the second group, mantle width exhibits
the greatest decrease, whereas siphon length the least
decrease, with the other characters intermediate. The less
rapid decrease mentioned above is not a constant decrease
as minor peaks are exhibited by head width at 40mm,
60mm, and 90mm; head length at 50mm; and nuchal
cartilage at 90mm.
Page 172
Fe)
aS
Morphometric Indices
°
iS)
ee
20 40 60 80 100 120 140 160
Dorsal Mantle Length
THE VELIGER
Vol. 12; No. 2
20 40 60 80 100 120 140 160 180
Dorsal Mantle Length
Figure 1
Relative growth patterns of Gonatopsis borealis (left) and Gonatus fabricu (right). The scales at the right of each are only for fin
width indices
FL Fin Length
FW Fin Width
HL Head Length
HW Head Width
Gonatus fabrici
The relative growth patterns of Gonatus fabriciu are
presented in Figure 1 (right). The two general patterns
observed for Gonatopsis borealis are also discernible for
this species. All morphological characters except fin length
exhibit growth curves that end at values lower than those
of their starting points. Most curves (MW, HW, HL, SW,
SL, and NCL) between 20 to 60mm mantle length show
minor but progressively lower fluctuations. Fin width and
fin length both show increases in the same interval, but
the curve for fin length continues to rise attaining a peak
at 143mm, and then decreases, whereas the curve for fin
width reaches a peak at 30mm and decreases thereafter.
DISCUSSION
Generally speaking, with two exceptions, the growth pat-
terns for Gonatopsis borealis and Gonatus fabricui are
similar, showing higher values in the smaller squid and
lower values in the larger squid. As the curves are of the
relative growth patterns (i. e., growth with respect to the
NCL Nuchal Cartilage Length
MW Mantle Width SL Siphon Length
SW _ Siphon Width
dorsal mantle length), this indicates that the mantle length
increases at rates greater than those of the other measure-
ments. As larval squid often have a truncated, broad
mantle and the mantle length: width ratios increase with
age, these patterns should be expected.
The two exceptions mentioned above are in fin width
and fin length. Both Gonatopsis borealis and Gonatus fab-
ric show large increases in the rate of growth for fin
length, but the rate of growth for the former reaches a
plateau after peaking at 30mm mantle length, whereas
the growth of the fin length of the latter continues to rise,
reaching a peak at 143mm mantle length. In the rate of
growth for fin width, Gonatopsis borealis exhibits rapid
growth, peaking at 50 mm mantle length, remaining steady
for another 40 mm of mantle length, then decreasing there-
after. The growth curve for fin width in Gonatus fabricu
increases rapidly to a peak at 30mm mantle length, but
then decreases throughout the rest of the curve. The
differences in fin morphology can be seen also in Figure 2.
I believe the above differences in the morphology of the
fins are indicative of a functional difference between the
two squid species. WILLIAMSON (1965) concluded from
Vol. 12; No. 2
observations made on Illex ilecebrosus that the fins served
to steer the squid and that, as the density of the squid
exceeded that of the water, they may also have served to
create a lifting force. Similar observations are mentioned
by Lane (1960) and Morton (1964). If the area of the
fin is related to lift (i. e., the larger surface area provides
greater lift when moving at slow speeds) then Gonatopsis
borealis (fin area = 478 mm?’ for dorsal mantle length of
Figure 2
External morphology of Gonatus fabricti (left) (217mm _ dorsal
mantle length) and Gonatopsis borealis (right) (210mm dorsal
mantle length)
210mm) would be more efficient than Gonatus fabricu
(fin area — 306mm? for a dorsal mantle length of 217
mm). The fin of Gonatopsis borealis is also stronger than
that of Gonatus fabrici, if greater fin thickness is indica-
tive of greater muscular strength. Non-quantified obser-
vations made during this study showed that there was a
difference in the thickness of the fins of the two squid
species.
THE VELIGER
Page 173
From the differences shown by the relative growth
patterns and by the gross external differences it appears
that the fins of Gonatapsis borealis play a much greater
role in the maintenance of vertical position and in loco-
motion at slow speeds than do those of Gonatus fabricii.
Although both squids are good swimmers, the former
should be more active than the latter in maintaining its
vertical position.
The relative growth patterns obtained in this study are
very similar to those calculated by HAEFNER (1964).
HAEFNER used the information from his morphometric
study as a means of taxonomically differentiating between
Loligo peale: and Lolliguncula brevis. He noted that the
main difference in the growth patterns of the two species
was in the fin width and length indices. Although he did
not reach any conclusions concerning the locomotion of
the two species, it seems as if functional differences hypo-
thesized in this study might also be applicable there.
ACKNOWLEDGMENTS
This work was done as a graduate student at Oregon State
University. I would like to thank Dr. Ivan Pratt of the
Department of Zoology for his assistance and for the lab-
oratory facilities. I would especially like to thank Dr.
William G. Pearcy, Department of Oceanography, for the
use of his laboratory facilities and for his suggestions
concerning this paper.
LITERATURE CITED
Haerner, Paut A,, Jr.
1964. | Morphometry of the common Atlantic squid Loligo
pealei, and the brief squid, Lolliguncula brevis, in Delaware
Bay. Chesapeake Sci. 5: 138 - 144
Lane, FRANK WALTER
1960. Kingdom of the Octopus.
York, N. Y.; 432 pp.
Morton, Joun Epwarp
1958. Molluscs: An introduction to their form and functions.
New York, Harper Bros. 232 pp.; 23 text figs.
Pearcy, WILLIAM GORDON
Sheridan House, New
1965. | Species composition and distribution of pelagic cephalo-
pods from the Pacific Ocean off Oregon. Pacif. Sci. 14:
261 - 266
WILLIAMSON, G. R.
1965. Underwater observations of the squid Illex illecebrosus
Lesueur in Newfoundland waters. Canad. Field-Natur.
79: 239 - 246
Page 174
THE VELIGER
Vol. 12; No. 2
Occurrence of the Sacoglossan Opisthobranch
Hermaea dendritica ALDER & Hancock in New England
BY
KERRY B. CLARK
AND
DAVID R. FRANZ
University of Connecticut Marine Research Laboratory, Noank, Connecticut 06320
THE PUBLISHED RECORDS of Sacoglossa from New Eng-
land include only six species: Elysia chlorotica (GouLp,
1870), Elysia catula (Gou.p, 1870), Alderia harvardiensis
GouLp, 1870 — considered to be synonymous with Alderia
modesta (LovEéN, 1844) (Hanp & STEINBERG, 1955; BLEAK-
NEY & BaiLey, 1967) — Stiliger fuscatus (GouLp, 1870),
Limapontia zonata (Grrarp, 1852), and Hermaea cruci-
ata Goutp, 1870. Of these, the latter 2 species have not
been reported since their original descriptions. Thus, a
new record would be the first addition to the fauna in
nearly 100 years.
Hermaea dendritica ALDER & Hancock, 1855, a species
occurring in Britain and Norway (Sars, 1878), the Medi-
terranean, and Japan (Marcus, 1961), has twice previ-
ously been reported from the western Atlantic. Marcus
(1961) reported the species from North Carolina, and
cited a manuscript by the late Dr. George Moore, noting
the occurrence of Hermaea dendritica in New England.
However, no description or analysis was given.
An established, breeding population of Hermaea dend-
ritica has been discovered near the Marine Research
Laboratory of the University of Connecticut, at Noank,
Connecticut, from which specimens have been collected in
May, 1967, and June to September, 1968. The specimens
agree closely with the description and figures of ALDER &
' Contribution No. 52 of the Marine Research Laboratory, Uni-
versity of Connecticut, Noank, Connecticut 06320.
Hancock (1846). Body color is cream-white or greenish,
with the liver ramifications visible through the body wall
as dark green or brown dendritic markings. The genital
opening is triaulic, and the albumen glands extend into
the cerata. There are from 18 to 65 smooth, club-shaped
cerata arranged in a double dorso-lateral row on each side.
The radula appears to be identical with that figured by
ALDER & Hancock (1846) and Bercu (1886).
Adults measuring 4 to 8mm in length have been
collected from the alga Codium fragile (SuRINGER) Hart-
oT 1867, which has only recently established itself in New
England. The Codium bed from which the specimens were
collected is located on a shell bottom in about 2.5 m of
water, at a salinity of about 30%,. Adults and egg masses
occur only on the upper, actively growing portions of the
algae. The egg masses contain 80 - 130 eggs measuring ap-
proximately 70 4, arranged in a broad flat coil of 1 - 14
turns, the entire mass measuring 1.0 to 1.6mm in dia-
meter.
Hermaea dendritica must be compared with two other
western Atlantic species, H. coitrala Marcus, 1961, from
Brazil (Marcus, 1961) and H. cruciata Goutp, 1870,
from Naushon Island, Massachusetts (Gouin, 1870).
Hermaea cruciata was briefly described by Goutp from
a sketch provided by Alexander Agassiz. No type specimen
of the species exists. ‘The sketch closely resembles. Marcus’
drawing of H. coirala. Both species differ from H. dendri-
tica in the following ways: the cerata are distally inflated,
Vol. 12; No. 2
with branching of the liver tubules at the tips, while those
of H. dentritica are of uniform thickness and the liver
tubules do not branch. The bodies of H. coirala and H.
cruciata are only half as broad as that of H. dendritica.
The oral veil of both species is markedly different, with
the anterior margin of the foot divided by a short groove
lacking in H. dendritica, and the cerata of H. coirala
and H. cruciata are arranged in two single rows. Hermaea
coirala’s rhinophores are distally bifid, but this feature
cannot be discerned from Agassiz’ sketch of H. cruciata.
Two factors imply that Hermaea dendritica has only
recently become established in the New England region.
First, as indicated by Gonor (1961), sacoglossans are
highly selective feeders, being restricted to one or a few
species of algae. Hence, it is highly unlikely that H. dend-
ritica could survive in New England until its food species,
Codium, had itself become established. Second, sampling
of many Codium beds in Eastern Connecticut has failed
to locate another population of the animal, a situation to
be expected where an organism has newly extended its
range.
THE VELIGER
Page 175
LITERATURE CITED
ALDER, JosHuA & ALBANY Hancock
1846. A monograph of the British nudibranchiate Mollusca.
Ray Soc., London
Bercu, Lupwic SopHus Rupo.tpH
1886. Beitraége zur Kenntniss der Aeolidaden, VIII. Verh.
Zool.-Bot. Gesellsch. Wien 35: 1 - 60
BLEAKNEY, J.S. & KANIAULONO BaILEy
1967. Rediscovery of the salt-marsh sacoglossan Alderia mo-
desta LovéN in eastern Canada. Proc. Malacol. Soc. Lon-
don 37: 347 - 349
Goutp, Aucustus ADDISON
1870. | Report on the Invertebrata of Massachusetts.
(Mollusca, W. G. Binney). Boston
Hanp, Capet & JOAN STEINBERG
1955. On the occurrence of the nudibranch Alderia modesta
Loven on the central Californian coast. The Nautilus
69 (1): 22-28
Marcus, ERNST
1961. | Opisthobranchia from North Carolina.
Elisha Mitchell Sci. Soc. 77 (2): 141-151
Sars, G. O.
1878. _ Bidrag til kundskaben om norges Arktiske fauna: I.
Mollusca regionis Arcticae Norvegicae. i- xii+466 pp., 34
+XVIII plts. Christiania
2nd ed,
Journ.
Page 176
THE VELIGER
Vol. 12; No. 2
Observations on the Tentacles of Vaginulus borellianus Couosi
(Mollusca : Gastropoda : Soleolifera )
BY
ARISTEO RENZONI
Istituto di Zoologia, Universita di Siena, Italia
(Plates 31 to 35; 3 Tables)
INTRODUCTION
THE TENTACLES OF GASTROPODS, like other organs of these
animals and of other metazoans, are able to regenerate.
In fact, it is quite certain that most of the deformities of
these cephalic structures are due to abnormal regeneration
following loss of or damage to tentacles. The first descrip-
tion we have of the phenomena of regeneration in mol-
lusks is that of SPALLANZANI (1768). Subsequently the
subject was taken up by various researchers (WarRTEL,
1768, ScHAFFER, 1768, MULLER, 1778, etc.) and analyzed
in great detail by Moguin-Tanpon (1851), who, in his
studies on Helix nemoralis, discovered that the regenera-
tion of the anterior portion of the head occurs only when
the cerebral ganglia remain intact. This finding explained
the seeming contradictions of previous researchers who
had cut off the tentacle or head at different levels.
Moguin-Tanpon’s results were confirmed by subsequent
studies on other gastropods (CarrizRE, 1880; CERNy,
1907; Konic, 1915). Nonne (1925) denies the importance
of the central nervous system in the regeneration of certain
parts of the tentacle, in particular the eye. ABELOOS
(1942) maintains that the regeneration of all cephalic
organs of gastropods is controlled by “a specific part of
the tegmentum.” After numerous histological and histo-
chemical studies of regenerating tentacles of Arion rufus
and Agriolimax agrestis, CHETAIL (1957, 1963) concludes
that after the removal of the tentacle a regeneration blas-
tema develops where the tentacle was amputated. It con-
sists of an exoblastema and an endoblastema, distinguish-
able by their different origins, histochemical character-
istics and systems of cellular reproduction.
None of the above mentioned authors has looked into
possible effects of the tentacles on the development or
functioning of the gonads. On the other hand, PELLUET
& Lane (1961) cut off the tentacles of Arion and Helix
with the main purpose of analyzing the effect on the
gonads. Among other interesting findings, they observed
that the optic nerve clearly regenerates after 3 to 4
weeks, and that, when only the optic and tentacular
nerves are severed, a great accumulation of secretion can
be found in the cytoplasm of the “collar cells” of the
tentacles.
My experiments with the amputation of tentacles were
carried out with the following purposes in mind:
1) To see whether regeneration occurs in Vaginulus
borellianus, and, if so, to analyse the phases and manner of
this process as well as the structure and the ultrastructure
of the regenerated organ;
2) To investigate the possible relationship between the
tentacle components (more precisely, their glandular and
neuroglandular elements) on the one hand and the devel-
opment of the gonads on the other.
Explanation of Plate 31
A) Dorsal section of the cephalic region of a normal animal. The
optic tentacles are retracted. 18
D) Same as Figure C.
yi = Be
PC = perigangliar cells
X75
OE
B) Sagittal section of an optic tentacle of a normal animal.
C) Distal end of the tentacular nerve. X30
x36
olfactory epithelium
TN = tentacular nerve
THE VELIGER, Vol. 12, No. 2 [Renzon1] Plate 31
Vol. 12; No. 2
THE VELIGER
Page 177
MATERIALS ann METHODS
Specimens of Vaginulus borellianus were obtained from
the Istituto di Zoologia, Universita di Firenze, about 3
years ago. They were kept in our laboratory in small
plastic boxes where they reproduced several times. Their
diet was lettuce, carrots, and slivers of fruit. For histo-
logical and histochemical studies the tentacles (both nor-
mal and regenerated) were fixed in Bouin’s fluid, neutral
10% formaldehyde or Susa’s fluid, dehydrated, embedded
in paraffin and sectioned, using the usual procedures. The
sections were stained with Mayer’s hemalum and eosin,
Heidenhain’s hematoxylin, PAS reaction (McManus),
and a modification of Bodian’s protargol-silver nitrate
method. For electron microscopy small pieces of normal
and regenerating tentacles were fixed in 3% glutaralde-
hyde in a 7.2 phosphate buffer, post-fixed in 1% osmium
tetroxide in a 7.2 buffer and embedded in Araldite. The
blocks were sectioned with an L. K. B. ultratome and the
sections examined in an Elmiskop I A electron microscope
after staining with uranyl acetate and lead citrate.
EXPERIMENTS
The following experiments were made:
A) amputation of the right optic tentacle of 10 10-day
old specimens;
B) amputation of the right optic tentacle of 8 6-month
old specimens ;
C) amputation of both upper tentacles of 6 10-day old
specimens ;
D) amputation of the lower tentacles of 6 10-day old
specimens ;
E) amputation of the upper tentacles of 8 10-day old
specimens followed by amputation of the regen-
eration blastema every 2 weeks over a period of
6 months;
F) amputation of both optic tentacles of 6 3-month old
specimens.
The tentacles were cut off under a binocular microscope
using partially anesthetized specimens with fully distended
tentacles. The regeneration blastema and control tentacles
were measured under the same conditions. In experiments
A and B, the left tentacle was considered the control
tentacle; for the other experiments 6 specimens of the
same age were used as controls.
OBSERVATIONS
Experiment A: The specimens were examined and meas-
ured 10, 30, 60, and 90 days after the amputation. The
measurements of the control tentacle, the regenerative
blastema and the regenerated tentacles thus obtained are
given in Table 1. It can be seen that regeneration occurs
at an even pace during the entire period of observation.
Two other facts seem equally noteworthy, namely that by
the 10" day, with the first phase of cicatrization completed,
the regeneration blastema is already evident and meas-
urable, and that some time between the 60" and 90" day
after the amputation, the regenerated tentacle has grown
to, or almost to, the length of the control tentacle. A
most interesting detail was found in one specimen of this
group. Serial sections showed that the regenerated nerve
consisted not of a single strand, as in the other regenerated
tentacles and the normal ones (Plate 31), but of 3
(Plate 32, Figure E), and that, at the tip of the tentacle
they end in the tentacular ganglion.
Experiment B: In this case too, all specimens were exam-
ined 10, 30, 60, and 90 days after the amputation; the
results are given in Table 2. Here, too, the rhythm of re-
generation is quite regular, and the regenerated tentacles
grow to the length of the controls within 90 to 100 days.
Experiment C: The specimens used in this experiment
were sacrificed at the age of 6 months and measurements
were made of the regenerated tentacles as well as the
hermaphrodite glands. Tentacles and hermaphrodite glands
of control specimens were also measured (see Table 3).
Experiment D: In this case the specimens were examined
Table 1
Regenerative Capacity of Tentacles of Vaginulus borellianus
Mean length in mm +standard error
L=length of left tentacle
R = length of right tentacle before and after amputation ( | )
Age in Days
10 20 30 40 70 100
L 2.07 +0.12 2.34+0.17 2.96 +0.14 4.27 £0.37 6.11+0.71 Weis 2040
R 2.01 +0.27 0.53+0.17 2.94 + 0.24 5.39+0.41 7.31+0.16
1.87 £0.09
Page 178 THE VELIGER Vol. 12; No. 2
Table 2
Regenerative Capacity of Tentacles of Vaginulus borellianus
Mean length in mm =standard error L=length of left tentacle
R = length of right tentacle before and after amputation ( | )
Age in Days
180 | 190 200 210 240 270
L 9.32 +£0.41 9.57 +£0.17 9.66 + 0.44 9.77 +£0.32 9.79 £0.22 9.80+0.21
R 9.47 £0.37 1.11+0.07 3.27+0.16 6.12 +0.37 8.96 £0.21 9.57 £0.39
Table 3
Weight of the body (b) ; mean weigth of the ovotestis (ot) (in gr)
and number of eggs (ne) in the ovotestis after amputation of
optic tentacles (O) or inferior tentacles (1) or repeated amputation
of the optic tentacles every 15 days (OO). C=Control individuals
Age in Days
10 30 60 90 120 150 180
O 0.094 0.201 0.473 1.023 2.633 4.300 5.98 +0.64 (b)
0.0246 + 0.0047 (ot)
130:0 =+1.98 (ne)
I 0.095 0.221 0.463 1.050 2.666 4.500 6.06 +0.77 (b)
0.0261 +0.0061 (ot)
WAS) sey (in)
OO 6.02 +0.49 (b)
0.0266 +0.0069 (ot)
28° Oe =17/7ieen (Ke)
Cc 0.066 0.202 0.471 1.039 2.550 4.450 3:83 027/ (by)
0.0288 + 0.0052 (ot)
133.0 +2.10 (ne)
Explanation of Plate 32
A) Section of a regenerating optic tentacle. X18 E) Dorsal section of a regenerating optic tentacle (Experiment A).
B) Same as Figure A at higher magnification. Note the numerous iihrce branches of the tentaculan TEtVe: zx 18
PEA he 60 F) Cross section of a regenerating tentacle (Experiment E). Arrows
Dee oe point to 4 branches of the tentacular nerve. 100
C) Perigangliar cells in a regenerating optic tentacle. 100 1 eidendnte GOS = gland of Semper’s organ
D) Regenerated lower tentacle. Layer of “Spindelzellen” (arrow) OE = olfactory epithelium PC = perigangliar cells
and gland of Semper’s organ. X 36 SpC = spindelzellen
TN = tentacular nerve
Explanation of Plate 33
Section of the distal portion of the olfactory epithelium of a normal
tentacle. X 30000
CG = cila D = dendrite M = mitochondria
ML = layer of mucus MV = microvilli
PP = plasmatic processes SC = sustentacular cells
SeC = sensory cells
[ReNnzon1] Plate 32
THE VELIGER, Vol. 12, No. 2
[RENzoNI] Plate 33
Tue VELIGER, Vol. 12, No. 2
Vol. 12; No. 2
THE VELIGER
Page 179
90 days after the amputation of the tentacles, at which
time the regenerated tentacles were of almost the same
length (5.94 mm) as those of the control specimens (6.05
mm). For the other results see Table 3 and Plate 32,
Figure D.
Experiment E: This experiment, in which the regenera-
tion blastema was re-amputated every 15 days, was de-
signed to eliminate any possible influence that the regen-
erating tentacle might have on the gonads. The blastema
developed again each time and the animal appeared to
be normal throughout the experiment. The results obtained
from 6 specimens are reported in Table 3. The other 2
specimens were kept alive for another 90 days after the
final amputation. By then the regenerated tentacles had
almost reached the length of those of the controls. In these
2 specimens the outward appearance of the regenerated
structure was variable; in general its diameter was greater
all along the tentacle and in particular at the tip, and
almost all the regenerated portions were colorless. Histo-
logical examination showed an almost total absence of
pigment cells in the epithelial and sub-epithelial layers.
In 2 specimens of this group the regenerated tentacular
nerves consisted not of a single strand, but of 4 to 5
branches (Plate 32, Figure F), closely resembling the
unusual specimen of group A. Since the material that was
fixed did not include the cerebral ganglia, I was not able
to trace these strands to their origin. However, we may
suppose that the branching of the tentacular nerve occurs
somewhere close to the cerebral ganglia.
Experiment F: The 6 individuals used in this experiment
were sacrificed 5, 15, 30, 60, 90, and 120 days after the
amputation of the tentacles, respectively. This was in or-
der to follow the stages in the differentiation of the
various components of the tentacles, as was recently done
with normal tentacles (RENZONI, 1968).
The following outline of events can be presented:
Until the 6" day, scar formation takes place with the
typical phenomena of pycnosis and the presence of numer-
ous macrophages of various sizes.
From the 7" to the 15" day, there is a period of latency
during which the appearance of the exoblastema and
endoblastema (typical regenerative processes) is clearly
seen.
From the 15" to the 40"™ day there is marked activity of
the components of the regenerating blastema. The large
number of different-sized nuclei indicates the presence of
the various cell populations. Mitotic figures are frequent
in the exoblastema, but rare in the endoblastema. Isolated
macrophages can still be found.
Between the 40" and the 90" to 100" day a gradual
return to normalcy takes place, following the differentia-
tion of the exo- and endoblastema into the various cell
types (Plate 32, Figures A, B) typical of the normal ten-
tacle: those of the distal sensory epithelium, of the tenta-
cular ganglion, the periganglion cells (Plate 32, Figure C)
(RENzONI, 1968), the “Spindelzellen” underneath the
epithelium (Plate 32, Figure B), the dermo-muscular
components, and the pigment cells branching among the
epithelial cells.
No instance of eye regeneration was found in this or any
of the other experiments. Some of the specimens were
observed directly under the dissecting microscope, but
most of them were sectioned serially. Neither method
showed any traces of eye formation, not even an invagi-
nation of the exoblastema that would have suggested a
possible retarded development of this organ. In fact,
several individuals from experiment A were kept alive
for more than a year and the eye still did not develop.
Regarding the ultrastructure of the components of the
tip of the regenerated tentacles, although no substantial
differences appear in the animals of experiments A and B,
they do appear in those of group E. The external appear-
ance and structure of the regenerated tentacle of animals
from this group showed considerable variation, and these
morphological changes were confirmed at the electron-
microscope, especially at the level of the sensory epi-
thelium. In fact, whereas the epithelium of the proximal
and medial portions of the tentacle, the “Spindelzellen”
and the few “collar cells” do not differ appreciably from
those of the normal tentacle, the cells that constitute the
olfactory epithelium are so changed (see Plates 33 and
34) that only by careful examination can one distinguish
the 3 cell types found in the sensory epithelium of normal
specimens (see RENZONI, 1968). ‘The sensory and the sup-
porting cells are particularly altered: in the former, the
the cytoplasm and ciliary apparatus are most changed; in
the latter, the apical microvillar processes (Plate 34; Plate
35, Figure A). The paucity of organelles (except for the
still numerous mitochondria) gives the cytoplasm of the
sensory cells the appearance of homogeneously low elect-
ron density. Some of the mitochondria are of the same
size as those found in the same elements of the normal
tentacle, some are much larger; however, at least with the
magnification used in this study, their structure does not
differ appreciably from that of the others. The microtu-
bules, which are especially numerous and evenly distrib-
uted in the sensory cells of the normal tentacle, are less
numerous and sometimes isolated and sometimes in clus-
ters (Plate 35, Figure C) in the regenerated tentacle.
Although the cilia at the apex of the dendrite have the
typical 9+2 pattern, their number varies considerably
from cell to cell. The microvilli also appear to be reduced
in number. The cytoplasm of the supporting cells is not
Page 180
THE VELIGER
Vol. 12; No. 2
noticeably changed, whereas the dense-cored filament of
the large microvillar processes of these elements under-
goes varying modifications (Plate 35, Figure D) ; in some
processes it is altogether lacking, in others it is present and
resembles normal ones, and in still others it is present, but
it is much larger than normal and of very high electron
density. Distally, the processes still have the tendency to
branch (Plate 35, Figure A) as in the normal tentacle,
and, furthermore, they seem to extend outwards to the
layer of mucus (usually covering the external surface of
the tentacles) with various projections of different sizes
and shapes (Plate 35, Figure A).
DISCUSSION
The process of regeneration of amputated tentacles (optic
and lower) in Vaginulus borellianus is substantially the
same as that described in the numerous studies on other
pulmonates.
The weights of the body and ovotestis and the number of
eggs in the ovotestis of experimental animals (experiments
C, D, E) show no significant variations either in compari-
son with each other or with the control animals. This is
in clear contrast with the results of PELLUET & LANE
(1961) in other pulmonates (Helix and Arion). These
authors find marked changes in the hermaphrodite gland
after the removal of tentacles and attribute them to the
absence of certain tentacular cells described as neurosec-
retory (hormonal). Morphologically similar cells, differ-
ing only in a few cytochemical and ultrastructural charac-
teristics, are also found in the periganglionic area of
tentacles of Vaginulus (RENZONI, 1968).
Regarding both structure and ultrastructure, whereas
the sensory cells in tentacles that have regenerated after a
single amputation do not differ appreciably from the con-
trols, those in tentacles that have regenerated after re-
peated amputations of the regenrative blastema are con-
siderably altered, especially at their apical end. Since, as
far as I can find, this is the first recorded instance of such
alterations, any hypothesis is risky, although it seems clear
that with the repetition of the trauma the nervous ele-
ments suffer most. The pigment cells are greatly dimin-
ished in experiment E, whereas the “Spindelzellen” do not
show appreciable differences.
In contrast with the results obtained by other authors
on different species, in our specimens of Vaginulus, whether
the tentacles were cut off once or the blastema was cut
off repeatedly, the eye consistently did not regenerate. The
results of the many studies of regeneration of organs and
structures in the cephalic region of mollusks, have always
shown discrepancies, sometimes because of the differences
in the species examined and other times because of mis-
taken interpretations. Nevertheless, the tendency has been
to attribute variations in regeneration to greater or lesser
damages suffered by the cerebral ganglia. In our study,
only the tentacles or regenerating blastema were cut off,
so that the eye’s incapacity to regenerate cannot be attrib-
uted to the removal of tissue from the cerebral ganglia.
For the present we are at a loss for an explanation of this
phenomenon; there is no doubt, however, that the non-
regeneration of the eye is further evidence that single or
Explanation of Plate 34
Distal portion of the olfactory epithelium of a regenerating tentacle
(Experiment E). 30000
ML = layer of mucus
Explanation of Plate 35
A) Same as the Figure of Plate 34. Note the distal branching of the
processes of the sustentacular cells. X 30000
B) Cross section of a dendrite of a sensory cell bearing cilia and
some microvilli. Regenerating tentacle. 30000
C) Cross section of a regenerating tentacle; microtubules in a
dendrite of a sensory cell. XX 30000
D) Cross section of the distal portion of a regenerating tentacle,
Where typical microvilli and a few plasmatic processes are
clearly evident. 30000
C = cilia ML = layer of mucus MT = microtubules
MV = microvilli PP = plasmatic processes
SC = sustentacular cells
THE VELIGER, Vol. 12, No. 2 [RENzonI] Plate 34
[RENzon1] Plate 35
THE VELIGER, Vol. 12, No. 2
Vol. 12; No. 2
THE VELIGER
Page 181
multiple amputations of the tentacle cause striking alter-
ations in components of the nervous system: branching of
the tentacle nerve, modification of the sensory cells, non-
regeneration of the eye.
SUMMARY
The author has conducted several experiments with the
amputation of the tentacles of Vaginulus borellianus
(Gastropoda Soleolifera) with the following purposes in
mind:
a) To see whether the regeneration occurs in this species
and if so, to analyse the phases and manner of this process
as well as the structure and ultrastructure of the regener-
ated organ;
b)to investigate the possible relationship between the
tentacle components (more precisely, their glandular and
neuroglandular components) on the one hand and the
development of the gonads on the other.
The following results have been obtained:
1) The process of regeneration on amputated tentacles
(optic and lower) in Vaginulus is substantially the same
as that described in the numerous studies of other pulmo-
nates.
2) The weights of the body and ovotestis and the number
of eggs in the ovotestis of experimental animals (experi-
ments C, D, E) show no significant variations either in
comparison with each other or with the control animals.
3) Regarding both structure and ultrastructure, whereas
the sensory cells in tentacles that have regenerated after a
single amputation do not differ appreciably from the con-
trols, those in tentacles that have regenerated after re-
peated amputations of regenerative blastema are con-
siderably altered, especially at their apical end.
4) The eye consistently did not regenerate in any of the
experimental animals (whether the tentacles were cut off
once or the blastema was cut off repeatedly).
LITERATURE CITED
Carriere, J.
1880. Studien tiber die Regenerationserscheinungen bei den
Wirbellosen. 1) Die Regeneration bei den Pulmonaten.
Wirzburg: 4-56 (as quoted by Cueralz)
Cerny, A.
1907. Versuche tiber Regeneration bei SiiBwasser- und Nackt-
schnecken. Arch. Entw. Mech. 23: 389 - 392
Cuneta, M.
1956. Caractéres histologiques du blastéme de régénération
tentaculaire chez Arion rufus L. Compt. Rend. Acad. Sci.
242: 1655 - 1656
1958. Action du bleu trypan sur le blasttme de régénération
tentaculaire d’Arion rufus L. Compt. Rend. Acad. Sci.
246: 642 - 644
1963. Etude de la régénération tu tentacule oculaire chez un
arionidae (Arion rufus L.) et un limacidae (Agriolimax ag-
restis L.). Ph. D. thesis, Facul. Sci. Univ. Paris. Masson &
Co., Publ.
Isserorr, H.
1964. Fine structure of the eye spot in the miracidium of
Philophthalmus megalurus. Journ. Parasit. 50 (4) : 549-554
Kernels, A.
1968. Nouvelles données histochimiques et ultrastructurales sur
les photorécepteurs “branchiaux” des Dasychone bombys
(DaLyELL) ; Annélide Polychéte. Zeitschr. Zellforsch. 86:
280 - 292
Konic, E.
1915. Die Regeneration des Auges bei Avion empiricorum.
Arch. Mikr. Anat. 80: 293 - 317
Mogurn-Tanpon, A.
1851. | Mémoire sur lorgane de l’odorat chez les Gastéropodes
terrestres et fluviatiles. Ann. Sci. nat. Zool. 15: 151 - 158
MULLER, M.
1778. Sur la reproduction des parties et nommément de la
téte des limagons a coquille. Journ. Phys. 12: 111 - 118
Nakamura, A.
1967. Electron microscopy on the ceras of an opisthobranch,
Godiva ceylonica, with special reference to muscles. Biol.
Journ. Okayama Univ. 13: 97 - 113
Nonneg, FE.
1925. | Versuche uber den Einflu8 des Nervensystems auf die
Regeneration der Augen von Pulmonaten. Arch. Entwickl.
Mech. 105: 430 - 469
PELLUuET, D. « N. Lane
1961. The relation between neurosecretion and cell differen-
tiation in the ovotestis of a slug. Gastropoda: Pulmonata.
Canad. Journ. Zool. 39: 789 - 805
RENZONI, ARISTEO
1968. | Osservazioni istologiche, istochimiche ed ultrastrutturali
sui tentacoli di Vaginulus borellianus (Colosi) ; Gastropoda:
Soleolifera. Zeitschr. Zellforsch. 87: 350 - 376
1968. Olfactory epithelium of gastropods. In: Electron
Microscopy (4 Europ. Region. Conf.) 2: 567 - 568. D. Steve
Bocciarelli, Ed.
ScuHArrer, M.
1768. | Versuche tiber die Reproduktion der Schnecken.
Regensburg.
SPALLANZANI, L.
1768. Prodromo di un’opera ad imprimersi sopra la ripro-
ductioni animali. Modena
Tonosaki, A.
1967. Fine structure of the retina in Haliotis discus. Zeit-
schr. Zellforsch. 79: 469 - 480
WartEL, C.
1768. | Mémoire sur les limagons terrestres de l’Artois pour
servir 4 l’histoire naturelle de cette province. Arras
Page 182
THE VELIGER
Vol. 12; No. 2
Marine Fouling and Boring Organisms in Monterey Harbor
II. Second Year of Investigation
BY
E. C. HADERLIE
Department of Oceanography, Naval Postgraduate School, Monterey, California 93940
(2 Text figures; 2 Tables)
INTRODUCTION
Since Octoser, 1966, continuous studies have been made
on the fouling and boring organisms that attach to or
drill into test panels exposed to the marine environment
under Municipal Wharf No. 2 in Monterey Harbor. The
results of the first year of study have been published
(HapERLIE, 1968a). The present paper will summarize
progress in the work during the second year, which basi-
cally extended from October, 1967, to October, 1968, but
continued through December, 1968, on certain exposed
panels. In addition to these investigations in the harbor
area, studies are also being conducted in the open water
of Monterey Bay in water-depths of 50, 100 and 200
feet. Results of two years’ work at the 50 foot level have
been published (Haperuiz, 1968b) and data from deeper
levels are presently being analyzed. It is planned to
continue all these investigations for several more years,
for experience has indicated that the results obtained
show great variability from year to year and data from
many years must be collected before a reasonable idea
of the nature of the fouling and boring community can
be obtained.
When this project was initiated in 1966 the objectives
were to obtain information on the kinds of marine organ-
isms that settle on or burrow into test panels of a variety
of types of material exposed in sea water at different
depths under the wharf, to determine the season or sea-
sons of settling, to note any correlation between settling of
organisms and the temperature or salinity or both of the
sea water, to determine any choice of substrate by indi-
vidual organisms, to measure rate of growth of the dom-
inant calcareous foulers, and to study evidence of seasonal
progression or ecological succession over an extended
period of time. Data from the first year’s work gave
partial answers to some of these problems, but experience
also indicated ways in which the experimental procedure
should be modified in order to get a more complete
picture. These modifications in technique will be explained
below.
The author wishes to acknowledge the following col-
leagues for help in the identifications of organisms: Mr.
Jack Gougé (Foraminiferans), Dr. D.J.Reish (Poly-
chaetes), Dr. D. P Abbott (Ascidians), Dr. A. H. Cheet-
ham (Bryozoans). Mr. Stephen V. Smith made miner-
alogical determinations of spirorbid tubes, and Mr. J. R.
Lance again assisted in resolving nomenclatural problems
with opisthobranchs. Mr. Barry Roth pointed out correct-
ly that the bivalves identified as Pecten sp. in my earlier
paper were in reality young specimens of Hinnites multi-
rugosus (GALE, 1928). Acknowledgment is also due Mr.
Jack C. Mellor for help in field work, to my wife, Mrs.
A. E. Haderlie, for assistance in the laboratory, and to the
Office of Naval Research for financial support.
AREA or STUDY
As reported in the 1968 paper on fouling and boring
organisms in the harbor, the site of the study is near the
outer end of Monterey Municipal Wharf No. 2 where
the water depth is approximately 21 feet at mean low
tide. For a complete description of the site readers are
referred to the earlier paper. As before, biweekly surface
temperature and salinity measurements were made
throughout the year (Figure 1).
METHODS
Techniques used during the period of study reported on
here were similar to those used the previous year, with
some modifications. During 1966-1967, collecting panels
consisting of standardized 8 inch by 10 inch panels of
Wolewli2INo: 2
THE VELIGER
Page 183
Temperature
Figure 1
Biweekly morning temperatures and salinities at test site for the year
October 1, 1967 to October 1, 1968
4 inch marine grade douglas fir plywood were used as the
primary collecting surfaces. Panels were placed in stain-
less steel racks holding 6 panels vertically and 3 inches
apart. To compare at least one other material that has
been used in the past as a collecting surface for foulers,
panels of 4 inch asbestos board (Johns-Manville Color-
lith) were also exposed in racks alongside those containing
plywood. It was found that there was essentially no dif
ference in the total number of species collected by ply-
wood or asbestos board and in settling most foulers did
not appear to distinguish between the two substrates. A
few organisms settled in slightly greater numbers on
plywood, whereas a few others were more abundant on
asbestos board, but the differences were not considered
significant. To simplify the procedure, during the 1967 to
1968-year reported on here the asbestos board panels
were discarded and only plywood panels used. Plywood
was selected over asbestos board, for wood panels also
collected wood boring organisms of interest in this study.
In the 1966-1967 study, one series of panels was sus-
pended in the water alongside the wharf where it was
exposed to relatively strong sunlight during at least half
Page 184
the day. These lighted panels collected a total of 5
species of benthic algae during the year, but otherwise
the fouling and boring organisms collected by the lighted
panels were very similar to those. collected by panels at
the same water depth in the dim-light conditions under
the wharf. During this past year, therefore, the lighted
series has been discontinued.
Again during the 1966-1967 year, one rack of panels
designated “floating panels” was rigged so that the rack
floated at the sea surface regardless of tidal level. This
rack was difficult to keep in place and several panels were
lost. During the 1967-1968 year the floating rack was
continually lost during storms and was replaced many
times. Ultimately this series of panels was abandoned.
This lack of data is perhaps not serious, for during 1966
to 1967 the floating panels did not collect a population of
foulers and borers much different from panels submerged
some distance below the surface.
During 1967-1968, then, three series of panels in racks
as used the previous year were continued as follows:
(1) Intertidal rack. Positioned about 4 feet above the
lowest low tide level. The panels in this rack were sub-
merged approximately one half the time and exposed
to air the other half. Short Term and Cumulative Panels.
(2) Shallow rack. Located 1 foot below lowest low tide
level. Panels always submerged. Short Term and Cumu-
lative Panels.
(3) Deep rack. Positioned 14 feet below the lowest low
tide level and about 7 feet off the bottom. Short Term
and Cumulative Panels.
In addition to the above listed three racks a fourth one
was added for reasons that will be explained below.
This rack was designated as follows:
(4) Shallow Long-Term Rack. Positioned 1 foot below the
lowest low tide level. Long Term Panels (3 month and
6 month exposure).
The basic period covered by the present study was
from October 1, 1967, to October 1, 1968. In the case of
the panels in the intertidal, shallow and deep racks the
routine for placing and retrieving the collecting surfaces
was the same as in 1966-1967. Of the 6 panels in each of
the racks there were 4 that were left in place for 3
months (Panel C-1), 6 months (Panel C-2), 9 months
(Panel C-3), and 12 months (Panel C-4) respectively.
These were designated Cumulative Panels. The other 2
panels in each rack were designated Short Term Panels
(S-1, S-2, etc.) : one was put in the water on the first of
each month and removed for analysis on the first of the
following month; the second panel was put in on the 15"
of each month and removed one month later.
THE VELIGER
Vol. 12; No. 2
During the 1966-1967 investigation it was found that
the Cumulative Panels exposed for 3, 6, 9, and 12 months
showed the most varied and extensive fouling communi-
ties. This was to be expected, for many benthic organisms
settle only on surfaces that have been colonized by pio-
neering fouling organisms. All of the Cumulative Panels
were placed in the water at the same time on October 1,
1966. During the course of the first year of this study it
was wondered if a panel placed in the water for 3 months
(or 6 months) beginning in October would ultimately
collect the same fouling community as one placed in the
water for 3 months (or 6 months) beginning in January,
March, or any other month. To find an answer to this
question a new rack was used during 1967-1968. The
rack was designated “shallow, long term rack” and was
positioned alongside the regular “shallow rack” at 1 foot
below low tide. Panels that would remain in the water
for 3 months (Panel 3-L) and for 6 months (Panel 6-L)
were submerged in this rack. One 3-month panel was
placed in the water on October 1, 1967, a second on No-
vember 1, 1967, and so on throughout the study period.
Six-month panels were submerged every 3 months. The
length of exposure of the various panels is shown diagram-
matically in Figure 2.
As before, the panels were removed from the racks
and transported to the laboratory in sea water containers
to be examined under a stereoscopic microscope while
the panels were submerged in a pan of sea water. After
examination, one of the surfaces of each of the Long Term
and Cumulative Panels (80 square inches or 5123 cm’)
was scraped clean of attached organisms and the scrapings
were oven dried at 100° C until the weight was constant.
This provided a rough statistical measure of the relative
amount of fouling growth accumulated in any one period
(see bottom line, Table 2). Short Term Panels usually
collected so little in terms of weight that they were not
scraped and treated as above.
THE FOULING COMMUNITY
I. DISCUSSION or ORGANISMS
SETTLING on SHORT TERM PANELS
Table 1 presents a list of organisms and their relative
numbers that settled on panels exposed for 1 month
during the period October 1, 1967, to October 15, 1968.
A total of 66 different animal species identified at least to
genus were recorded. During the previous year, a total of
70 different kinds of organisms was found on similar
panels, but this number included 4 species of algae. None
of the panels submerged under the wharf during 1967
Woltal 2 No: 2
THE VELIGER
Page 185
Interval of Exposure
ANNNNUNANNADN
Or Ou fw hd we
n
°
Short Term Panels
(3 months)
Long Term Panels
(6 months)
Cumulative
aE ee
Pf ffs [Fp [mis [7 Jas Jol] >
Figure 2
Diagram illustrating period of exposure for test panels from
October 1, 1967 to January 1, 1969
to 1968 collected any green, brown, or red algae for, as
explained above, none of the racks was placed in water
receiving direct sunlight. All panels when first placed in
the water collected a surface film of bacteria and benthic
diatoms, but none of these was identified.
During each of the past 2 years, then, Short Term
Panels have collected 66 different kinds of animals. This
identical total seems to be merely a coincidence for, al-
though the totals were the same, the lists of organisms
are not identical for each of the two years. During each
year the dominant fouling organisms were the same: en-
crusting cheilostomate and cyclostomate bryozoans, cal-
careous-tubed serpulid worms and acorn barnacles, The
most common boring organism was the gribble Limnoria,
and these and other wood borers will be considered later.
The following discussion will briefly review the occurrence
of the most common organisms of each major group listed
in Table 1.
Protozoa:
Eight different kinds of benthic foraminiferans were
observed on the Short ‘Term Panels. Cornuspira lajollensis
and Rosalina columbiensis were the only species regularly
encountered, but the former was often found in great
numbers. The reason that more species of foraminiferans
were found during 1967-1968 than in the previous year
was due in part to a more determined effort to locate
them, but principally because Mr. Jack Gougé examined
many of the panels when fresh and picked up forms that
had been overlooked previously. The ciliate Folliculina sp.
was perhaps the most abundant single protozoan encoun-
tered, and it settled on panels at all depths throughout
the year. The suctorian Ephelota gemmipara often formed
fuzzy forests over the panel surface and attached to hydro-
ids such as Obelia. The colonial ciliate Zoothamnium sp.
again was found to be common during the summer and
fall months.
Porifera:
Two species of sponges settled on the Short Term Pan-
els, but of these only Leucosolenia eleanor was found
regularly.
Coelenterata:
The only coelenterate that settled on the panels with
any regularity was the hydroid Obelia sp. which settled
during most months of the year. Syncoryne mirabilis, a
second hydroid, was encountered only rarely and in small
numbers.
Page 186
THE VELIGER
Vol. 12; No. 2
Platyhelminthes:
Four different species of flatworms were recorded on
Short Term Panels during 1967-1968. The commonest
form encountered was Thysanozoon californicum which
occurred only on those panels with a population of the
bryozoan Celleporaria brunnea.
Nemertea:
Nemertean worms were rarely seen on the panels, but
two species were observed, each on one occasion, on the
intertidal panels only.
Ectoprocta (Bryozoa):
{Dr. Cheetham of the U.S. National Museum has examined
the cheilostomate bryozoans on the panels and has found that
the bryozoan identified as Lyrula hippocrepis (Hincks, 1882)
in my earlier papers (HapErR.iE, 1968a, 1968b) is in reality
Cryptosula pallasiana. In addition, Hippothoa hyalina should
be called Celleporella hyalina and Holoporella brunnea should
be called Celleporaria brunnea.}
During the present study the same 4 encrusting bryo-
zoans dominated the Short Term Panels as they did in
1966-1967. Celleporella hyalina, Cryptosula pallasiana,
Tubulipora pacifica and Celleporaria brunnea were found
on nearly all panels at all depths. During this past year
Celleporaria seemed to be much more abundant, and
Celleporella less abundant than during the year before.
Erect bryozoans were also very common. These were dom-
inated by the soft fuzzy Bowerbankia gracilis and by
Bugula neritina and B. californica. On a shallow panel
submerged from June 15 to July 15, 1968, dozens of tiny
Bugula neritina were found, indicating the period of maxi-
mum settlement. During most months of the year only
one or two specimens of B. neritina were found on any
panel. From mid-summer through the fall Crisulipora
occidentalis settled on panels in the shallow and deep
racks.
Annelida:
During 1966-1967 spirorbid worms were encountered
on the Short Term Panels in abundance second only to
encrusting bryozoans. These forms were identified as Spir-
orbis spirillum and they settled on the panels at all depths
throughout the year. During 1967-1968 spirorbids were
again encountered, often in great numbers, and on most
Short Term Panels throughout the year. The most intense
settling was from mid-April through June when hundreds
of small worm tubes were found, particularly on the
deep panels. A more careful examination of the tubes of
these spirorbids has shown that at least 4 distinct morpho-
logical types are present. The largest tubes and most
commonly encountered forms are distinctly sculptured,
coiled counter-clockwise, and faintly resemble the tubes of
Spirorbis nipponicus Oxupa, 1934, as figured in USHakov
(1955). Mineralogical analysis of the tubes by Stephen V.
Smith has shown these to be composed of aragonite. A
second morphological type (and second in abundance on
the panels) coils clockwise and has a tube strongly
ringed resembling the horn of an antelope such as the
sable or oryx. These tubes are composed of calcite with
12% Mg substitution for Ca in the CaCO: lattice. A
third form has a very smooth shell and coils clockwise.
It is composed of aragonite. The fourth form was only
rarely encountered. It is smooth and coils counter-clock-
wise and also seems to be composed of aragonite. When
removed from the tubes the worms all look remarkably
alike. The identity of these spirorbids is therefore in
doubt. It is possible that these forms represent 3 or 4
distinct species. Until the systematics is worked out it
seems wise to refer to them as Spirorbis spp.
A second common serpulid that has been found not
only in Monterey Harbor but at depths to 100 feet in
open water has been referred to previously only as Serpula
sp. (HapEruE, 1968a, 1968b). A more critical examina-
tion of these worms during recent months has shown them
to be Chitinopoma occidentalis. On deeper panels Chitin-
opoma was more abundant than S‘prrorbis.
Arthropoda:
As in the earlier study in the harbor, the most common-
ly encountered arthropod on the fouling panels was the
acorn barnacle Balanus crenatus. On the Short Term
Panels this barnacle settled in small numbers during most
months of the year. There were usually 10 animals or
fewer per panel side, and the greatest number seen was
20 per panel side during August, 1968. These results ob-
tained during the 1967-1968 year contrast sharply with
Explanation to Table 1 (foldout, facing this page —>)
* Symbols used at head of columns indicate:
S-1, S-2, etc. = Short term panel number as designated in
Figure 2
I = Intertidal panels
S = Shallow panels
D = Deep panels
2 Symbols used in columns indicate:
1 = species present in numbers from 1 to ro individuals or
colonies per panel side
2 = species present in numbers from 11 to 20 individuals or
colonies per panel side
3 = species present in numbers upward from 20 individuals or
colonies per panel side
*
ui.
pari
Site Miles nee
List of Species, and of Panels where Organisms Settled in Short Term Series
Table 1
Protozoa:
: Jridia lucida LECALUEY, 1936
Cornuspira lojollaensis Uctito, 1960
Rosalina columbiensis (Cusnman, 1925)
Spirillina revertens (Ruumster, 1906)
Spirillina vivipara (RHUMBLER, 1906)
Patellina corrugata Wriiiamson, 1858
Tubinella funalis (BRADY, 1884)
Entosolenia sp.
Fildile gemmipara (Hartwic, 1876)
Zoothamnium sp.
Folliculina sp.
Stentor sp.
Porifera:
Leuconia heathi (Urpan, 1905)
Leucosolenia eleanor URBAN, 1905
<
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S-12
S|D
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Coelenterata:
Obelia sp.
Syncoryne mirabilis (AGASSIZ, 1862)
Metridium senile (LinNapus, 1767)
Platyhelminthes:
Stylochoplana gracilis HeatH & McGrecor, 1912
Notoplana acticola (Boone, 1929)
Thysanozoon californicum Hyman, 1953
Pseudocerous sp.
NO DATA
— NO DATA —
=
Nemertea:
Tubulanus sexliniatus (Grirrin, 1898)
Tetrastemma nigrifrons Cor, 1904
N. D.
Entoprocta:
Barentsia gracilis (Lomax, 1886)
N. D.
Ectoprocta (Bryozoa) :
Bowerbankia gracilis O'Donoonue, 1926
Filicrisia franciscana (RobERTSON, 1910)
Crista sp.
Crisulipora occidentalis Ropertson, 1910
Tubulipora pacifica Robertson, 1910
Bugula neritina (Linnazus, 1758)
Bugula californica Ronertson, 1905
Celleporella hyalina (LinnNazus, 1767)
Cryplosula pallasiana (Mott, 1803)
Scrupocellaria californica Tras, 1857
Scruparia ambigua (v'Orbiony, 1841)
Celleporaria brunnea (HincKs, 1884)
NO DATA
new
enn
Annelida:
Platynereis agassizi (EHLERS, 1868)
Pseudopotamilla ocellata Moore, 1905
Spirorbis spp.
Chitinopoma occidentalis (Busu, 1904)
— NO DATA —
=EE Ho
Po Ce
palin colar,
IR
Sa
ical
n
3
Arthropoda;
Balanus crenatus Bruouttre, 1789
Balanus glandula Darwin, 1854
Balanus tintinnabulum (Linnagus, 1758)
Chthamalus dalli Pruspry, 1916
Lepas anatifera (Linnazus, 1758)
Caprella californica Stimpson, 1857
Corophium insidiosum Crawrorp, 1937
Limnoria quadripunctata Hornuts, 1949
NO DATA
Loxorhynchus crispatus Stimpson, 1875
Mollusca:
Pododesmus cepio (Gray, 1850)
Hinnites multirugosus (Gate, 1928)
Mytilus edulis Linnazus, 1758
Hiatella arctica (LINNAEUS, 1771)
Bankia setacea (Tron, 1863)
Coryphella trilineata O'Donoanue, 1921
Hermissenda crassicornis (Escuscnoxtz, 183!)
Dendronotus frondosus (Ascantus, 1774)
Dendronotus subramosus MacFARLAND, 1966
Doto kya Marcus, 1961
Tiiopha grandis MacFartanp, 1905
Trinchesia albocrusta (MacFartanp, 1966)
Polycera atra MacFARLAND, 1905
NO DATA
: Aegires albopunctatus MacFarLanp, 1905
Kehinodenmata:
Strongylocentrotus sp.
—CPhiothrix Spiculata LeConte, 1851
NO DATA\
[os
Vol. 12; No. 2
THE VELIGER
Page 187
the former year where during certain months such as
March, June and August there were massive setlements
of up to 25 barnacles per square inch of panel surface.
During the present study there was no one period when
B. crenatus settled in great numbers. On panels in deeper
water in the open bay, where massive settlement of B.
crenatus occurred in March and August, 1967, no such
peak in settling was recorded in the same months of
1968 (Haper.iz, 1968b). Thus the data from the harbor
and from open water are in agreement; no major seasonal
settlement of B. crenatus occurred in 1968. The reason
for this is not obvious. The water temperature and salin-
ity recorded in 1968 was not much different from the
previous year (Figure 1). This shows again that one
must make observations over a number of years before one
can generalize regarding season of settling of fouling
organisms.
The small acorn barnacle Chthamalus dalli was re-
corded fairly regularly from panels exposed in the inter-
tidal rack. The peak period of settlement was between
February 15 and March 15, 1968, when over 100 barnacles
settled on each side of a wooden intertidal panel. It was
interesting to note that the barnacles settled only on the
dark areas of the wood and on pencilled letters used
for marking the panels. Perhaps a very dark board or a
panel of asbestos board would have collected many more
of these.
Other barnacles recorded from Short Term Panels were
a few specimens of Balanus glandulus, one of B. tintin-
nabulum and one of Lepas anatifera.
Mollusca:
The nestling clam Hiatella arctica was commonly found
on Short Term Panels as a tiny, freely-moving organism.
In the spring months there were often dozens of these on
every panel. Pododesmus cepio was also found fairly regu-
larly throughout the year, but only in small numbers.
Mytilus edulis, a dominant animal on the pilings of the
Wharf, settled only occasionally on Short Term Panels.
Many nudibranchs were found but of these only Hermis-
senda crassicornis was of regular occurrence.
Echinodermata:
Small green sea urchins were recorded on most panels
during the spring and summer months. These were invari-
ably so small as to make specific identification impossible.
Chordata (Tunicata):
As was true in the previous year, during 1967-1968 no
tunicates settled on the Short Term Panels, yet, as will be
seen, did settle on the Long Term and Cumulative Panels.
This again confirms what ScHEER (1945) and others
have found, namely, that tunicates settle only on surfaces
that have been colonized by earlier foulers such as
bryozoans.
II, DEPTH PREFERENCE
As can be seen from Table 1, most of the organisms
encountered in this survey showed little preference for
panels at different depths, and any one organism could be
found on panels at all depths at one time or another.
Exceptions to this generalization, however, were found
among certain of the encrusting bryozoa. Tubulipora pa-
cifica, for example, was exceedingly common on the con-
tinually submerged panels, but was found on only one
occasion on an intertidal panel. Celleporaria brunnea was
also common on submerged panels, especially the shallow
panels, but rare on the intertidal ones. Crisulipora occi-
dentalis definitely preferred the deep panels. On the other
hand, the barnacles Balanus glandula and Chthamalus
dalli were found only on intertidal panels.
III. DISCUSSION or ORGANISMS
SETTLING on LONG TERM anno CUMULATIVE
PANELS
Table 2 presents data collected from Long Term and Cu-
mulative Panels during 1967-1968. As indicated in Figure
2 and discussed earlier in this paper, the panels desig-
nated Long Term Panels were those that were immersed
in racks suspended one foot below low tide level. Most
of these panels were exposed for 3 month periods, one
being submerged on the first of every month throughout
the year and removed for analysis 3 months later. A
second group of 4 Long Term Panels in the same rack
remained in the water for 6 months; one panel was
immersed on October 1, a second on January 1, a third
on April 1 and the fourth on July 1.
Cumulative Panels, on the other hand, were submerged
at the 3 depths discussed earlier in connection with Short
Term Panels, namely in intertidal, shallow and deep
racks. All the Cumulative Panels were submerged at the
same time on October 1, 1967. From each of 3 racks one
panel was removed at the end of 3 months, 6 months, 9
months and finally at the end of 12 months. The data
from these panels are given in Table 2, right side. The rea-
son there is no column for data from shallow panels at 3
and 6 months is that these data were incorporated into the
Page 188
THE VELIGER
Vol. 12; No. 2
appropriate 3 or 6 month columns under Long Term
Panels.
On these Long Term and Cumulative Panels a total
of 88 different animals identified at least to genus were
recorded. This was about the same total number as
found in 1966-1967, but the list does include several
animals not recorded earlier, particularly foraminiferans.
And because none of the panels was exposed to bright
light, no benthic algae were found on the 1967-1968 series.
In the discussion that follows, the dominant animals of
each major group that settled in any numbers on the Long
Term and Cumulative Panels will be discussed briefly.
Protozoa:
A total of 15 different kinds of benthic foraminiferans
were recorded during the year, Many of these were found
on only one or two occasions, but Cornuspira lajollensis
was found regularly throughout the year and often was
present in great numbers.
The ciliate Folliculina sp. was also exceedingly common
throughout most of the year. On Cumulative Panels in
the deep rack removed after 3, 6, 9, and 12 months, this
large ciliate was often one of the dominant foulers in
terms of numbers of animals present. On the deep panel
removed on January 1, 1968, after 3 months exposure,
Folliculina was present in thousands and blackened the
surface of the panel. The same was true of a shallow panel
removed April 1, 1968.
Porifera:
The sponge Leucosolenia eleanor was encountered on
nearly all of the panels except the Cumulative Panels
exposed in the intertidal. The anastomosing tubes often
formed large clusters 5 cm or more in diameter on panels
in the water for 3 months or more. Large globular speci-
mens of Leuconia heathi up to 3cm in diameter were
present in fair numbers on deep panels in the water for 6
months, and on the 9-month panel it was second only to
the bryozoan Crisulipora occidentalis as the dominant
organism.
Coelenterata:
No coelenterates were common on the panels. Obelia
sp. was found sporadically, but only on an intertidal panel
in the water 6 months was it ever abundant. Metridium
senile was recorded only twice: on an intertidal and a
deep panel exposed 3 months beginning October 1, 1967.
On each panel there was a single anemone | cm in dia-
meter. It is surprising that more individuals of Metridium
did not settle on the panels, for this anemone along with
Corynactis californica CaRLGREN, 1936, is among the
dominant fouling organisms on the wharf pilings near the
position where the collecting panels were exposed. It is
possible that a new surface must be in the water for a
period much greater than 12 months before it is suitable
for settlement of these anemones. In the 1966-1967 survey,
Corynactis was encountered a few times on panels in the
water 9 months or more, but this past year none were
recorded.
Platyhelminthes:
Six species of flatworms were found on the Long Term
and Cumulative Panels but of these only 'Thysanozoon
californicum was common. This very unusual flatworm
is papillated on the dorsal surface and in color and texture
closely matches the encrusting bryozoan Celleporaria
brunnea on which it lives. The worm is elliptical in out-
line and the largest individuals seen were 10 mm long.
Thysanozoon was encountered regularly, especially on the
3 and 6 month Long Term Panels, and throughout the
year except for late spring and early summer. Never was
Thysanozoon found on a panel unless Celleporaria was
established there, and in all cases the flatworm was found
pressed tightly against the bryozoan colony. It was found
to be very sluggish and moved only when prodded. When
the flatworm was loosened and turned over it was often
seen to have the pharynx extended and pressed over the
zooids and in many cases the zooecium had turned white
in color and the living zooids were gone. This indicates
strongly that Thysanozoon feeds directly on the zooids of
Celleporaria.
Ectoprocta (Bryozoa) :
As can be seen in Table 2, the bryozoans were clearly
the dominant fouling animals on the Long Term and Cu-
mulative panels during the 1967-1968 year as they had
Explanation to Table 2 (foldout, facing this page —>)
* Symbols used at head of columns indicate:
3L1, 3L2, ... 6L1, 6L2, etc. = Panel numbers as designated
in Figure 2
I = Intertidal panels GC
S = Shallow panels
D = Deep panels
2 Symbols used in columns indicate: .
I = species present in numbers from 1 to 10 individuals
or colonies per panel side
2 = species present in numbers from 11 to 20 individuals
or colonies per panel side
3 = species present in numbers upward from 20 individuals
or colonies per panel side
VEC INOS
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A bose ie ea ncn tenn sete ide ays pte by ear yest) AS | pix 1 aoa (6),
ack
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Table 2
List of Species, and of Panels where Organisms Settled in Long Term and Cumulative Series, October 1, 1967 to January 1, 1969
Species
Protozoa:
Inidia lucida LeCarury, 1936
Cornuspira lajollaensts Ucuto, 1960
Rosalina columbiensis (Cusuman, 1925)
Spirillina revertens (RHuUMBLER, 1906)
Spirillina vivipara (RHUMBLER, 1906)
Tubinella funalis (Bravy, 1884)
Saccamina sp.
Cibicides sp.
Rotorbinella campanulata (GALLOWAY & WISSLER, 1927)
Haplophragmoides columbiense CusuMan, 1925
Quinqueloculina sp.
Spiroloculina hyalina Scuuze, 1875
Miliolinella sp.
Bolvina sp.
Gordiospira sp.
Folliculina sp.
Zoothamnium sp.
Ephelota gemmipara (Hartwic, 1876)
Long Term Panels
Porifera:
Leuconia heathi (Urpan, 1905)
Leucosolenia eleanor Urnan, 1905
Halichondria panicea (Pautas, 1766)
Rhabdodermella nuttingi Ursan, 1902
Coelenterata:
Obelia sp.
Syncoryne mirabilis (Acassiz, 1862)
Metridium senile (Linnazus, 1767)
Platyhelminthes:
Kaburakia excelsa Bock, 1925
Stylochoplana gracilis Heatu & McGrecor, 1912
Notoplana acticola (Boone, 1929)
Thysanozoon californicum Hyman, 1953
Stylochus sp.
Pseudocerous sp.
Nemertea:
Tubulanus sexliniatus (GrirFin, 1898)
Micrura verrilli Cor, 1901
Amphiporus bimaculatus Cor, 1901
Amphiporus imparispinosus Grirrin, 1898
Entoprocta;
Barentsia gracilis (Lomax, 1886)
Ectoprocta (Bryozoa) :
Bowerbankia gracilis O’Donocuur, 1926
Filicrisia franciscana (Ropertrson, 1910)
Crisia sp.
Crisulipora occidentalis Rosrrtson, 1910
Tubulipora pacifica Roverrson, 1910
Bugula neritina (Linnagus, 1758)
Bugula californica Ronertson, 1905
Celleporella hyalina (Linnaeus, 1767)
Cryptosula pallasiana (Mout, 1803)
Scrupocellaria californica Trask, 1857
Scruparia ambigua (p’Oruicny, 1841)
Celleporaria brunnea (Hincxs, 1884)
Dendrobeania lichenoides (Ronertson, 1900)
Annelida:
Platynereis agassizt (Euvers, 1868)
Nereis vexillosa Grune, 1851
Halosydna brevisetosa Kinnerc, 1855
Sabellaria cementarium Moore, 1906
Sabella sp.
Pseudopotamilla ocellata Moore, 1905
Spirorbis spp.
Chitinopoma occidentalis (Busu, 1904)
Polydora brachycephala Harrman, 1936
Sipunculoidea:
Phascolosoma agassizit KEFERSTEIN, 1866
Arthropoda:
Balanus nubilis Darwin, 1854
Balanus crenatus Brucuitre, 1789
Balanus glandula Darwin, 1854
Balanus tintinnabulum (LINNaA®’
Chthamalus dalli Prrsnry, 1916
Caprella californica Stimeson, 1857
Corophium insidiosum Grawrorp, 1937
Limnoria quadripunctata Houruurs, 1949
Loxorhynchus_crispatus Sviaeson, 1875
Mollusca:
Pododesmus cepio (Gray, 1850)
Hinnites multirugosus (Gaus, 1928)
Mytilus edulis Linnasus, 1758
Hiatella arctica (Linnagus, 1771)
Bankia setacea (TRyon, 1863)
Hermissenda crassicornis (EscuscHoxrz, 1831)
Acanthodoris brunnea MacFarvanp, 1905
Doto kya Marcus, 1961
Triopha carpenteri (Srearns, 1873)
Triopha grandis MacFartanp, 1905
Trinchesia albocrusta (MaGFartanp, 1966)
Polycera atra MacFartann, 1905
Aegires albopunctatus MacFartanp, 1905
Echinodermata:
Strongylocentrotus purpuratus (Stinson, 1857)
Strongylocentrotus franciscanus (Agassiz, 1872)
Strongylocentrotus spp.
Ophiothrix spiculata LyConts, 1851
Rupentacta quinguesemita (Seuenxa, 1867)
Chordata (Tanicata) :
Aplidium solidum (Rrrrex & Forsytx, 1917)
Ascidia ceretodes (HUNTSMAN, 1912)
Styela truncata Rerrsr, 1901
Dry weight (gr) of fouling growth
s, 1758)
9.2
9.0) 8.1
7.3)
10.0)16.1]
13.0}22.41
10
3L
12
3L
6L
Cumulative Panels
6 months 6 mos.
1| 2) 3
6L |6L S
3 mos. 9 mos.
eee
1513
19.2/13.5 6.8
me
|
|
25.1/61.3/29.7} 1.0} 5.0] 2.3/14.7|30.6/77.6}45.94 0.0/109 32.3
meee
Vol. 12; No. 2
THE VELIGER
Page 189
been the previous year. Twelve species were found regu-
larly, and often in great numbers, and many of these
settled on panels immersed at different periods during the
year indicating that there are settling larvae present
througout the year. In many cases it was difficult to decide
which of these bryozoans were truly dominant, for in
many cases a panel was completely covered with encrust-
ing colonies of Tubulipora pacifica, Cryptosula pallasiana
and Celleporaria brunnea. In other cases the panel might
be practically covered with a thick growth of Crisulipora
occidentalis sprinkled with fewer colonies of Bugula neri-
tina and B. californica.
On the Cumulative Panels in the water 3, 6, 9, and 12
months from October 1, 1967, it was obvious that most of
the bryozoans preferred the panels in the shallow and
deep racks where they were constantly submerged. Only
Bowerbankia gracilis and Cryptosula pallasiana occurred
commonly and abundantly on intertidal panels, and on
the 9 and 12 month intertidal panels Bowerbankia was the
dominant animal and formed a fuzzy mass over every-
thing. Some of the other forms also showed a depth pref-
erence. Crisulipora occidentalis, for example, was found
commonly on panels at both the shallow and deep posi-
tions, but only on the deep ones did it become the domi-
nant form. On panels in the shallow position Celleporaria
was most abundant with Cryptosula and Tubulipora next.
As regards the Long Term Panels that were exposed in
racks at the shallow depth for 3 or 6 months but at
varying times throughout the year, the dominant animal
in all cases was Celleporaria brunnea. Every panel of this
series collected massive numbers of Celleporaria colonies.
At the end of any 3 month period, the wooden panel
would be practically totally encrusted with these colonies,
each averaging 10 to 20 mm in diameter, with Cryptosula
and Tubulipora filling any blank space. Often the Celle-
poraria colonies had completely overgrown and smothered
other encrusting bryozoans such as Celleporella hyalina,
acorn barnacles and serpulid worm tubes. It is interesting
to note also that Celleporaria settled fairly evenly and had
similar growth rates throughout the year, although most
of the colonies were slightly smaller in size in the winter
months compared to the summer. In other words, a
panel removed in January, in May, in August or in Octo-
ber after being in the water 3 months had a very similar
population of Celleporaria. This, however, held for panels
in the shallow position only. No equivalent series was
exposed in deeper water under the wharf, but circum-
stantial evidence from other deep panels would indicate
that perhaps Crisulipora occidentalis would be the dom-
inant form throughout the year at this position.
Annelida:
Calcareous serpulid worms were commonly found on
most Long Term and Cumulative Panels. The most abun-
dant of these were spirorbids which will be simply referred
to as Spirorbis spp. A description of the 4 morphological
types of these spirorbids, based on the tube form and
shape, was given earlier under the section dealing with
annelids on Short Term Panels. On the Long Term Panels
the largest and most abundant of these were those with
sculptured tubes spiralling counter-clockwise. Many of
these were 2.5 mm in maximum dimension. The smallest
were the smooth forms spiralling counter-clockwise, the
largest of which were 0.9 mm across.
The second common serpulid was Chitinopoma occi-
dentalis (referred to incorrectly in 2 earlier papers as
Serpula sp. (HaperuE, 1968a, 1968b) ). This serpulid is
distinct in having a tube with a prominent keel running
its full length and ending in a denticle over the orifice.
The tubes were often twisted and distorted, particularly
on older panels and measured up to 50 mm in length.
Arthropoda:
During the 1966-1967 survey and in earlier pilot
studies in the harbor (MomMsEN, 1966; Miter, 1966),
the acorn barnacle Balanus crenatus was one of the com-
mon foulers on Cumulative Panels and was often the
dominant animal. As pointed out in the discussion of B.
crenatus on Short Term Panels, the 1967-1968 season
showed very few settling barnacles anywhere in the har-
bor or out in open water of the bay. It is not surprising,
then, that Long Term and Cumulative Panels collected
few barnacles. At no time were more than 20 B. crenatus
found on any one panel, regardless of the length of
exposure.
Balanus glandula and Chthamalus dalli occurred in
small numbers on a few of the intertidal Long Term
Panels, and Balanus tintinnabulum on a few intertidal
and shallow panels. The giant acorn barnacle, Balanus
nubilis, made a single solitary appearance on an inter-
tidal panel exposed for 9 months. This barnacle which
is exceedingly common on the piles in Monterey Harbor
has not been recorded from a test panel before during
these studies.
Mollusca:
A variety of mollusks was encountered on the Long
Term and Cumulative Panels, most of them opistho-
branchs, and usually in small numbers. Hermuissenda
crassicornis was the most common of these. The nestling
Page 190
clam Hiatella arctica was fairly common on panels that
had an extensive growth of erect bryozoans or sponges
that created crevices in which this animal could live. On
a shallow panel exposed 3 months and removed April 1,
1968, there were hundreds of small Hiatella nestled in the
bryozoans.
Mytilus edulis is a very common fouler on the wharf
pilings of the harbor, yet it settled only in small numbers
and then only on panels in the water 6 months or more
and in the intertidal position. The maximum size was
4 cm and this was achieved on a panel exposed 12 months.
Yet on the intertidal rack itself a large cluster of about 30
individuals of M. edulis was found attached to the board
that formed the bottom of the rack. Some of these were
large (7 cm and more). As mentioned above, no mussel
was found on any panels exposed for less than 6 months,
so it is possible these large animals had developed in
half a year or less.
Echinodermata:
Small sea urchins (Strongylocentrotus spp.) 2.0mm
and less in diameter were found on most of the Long
Term Panels and occasionally in fairly great numbers.
These were impossible to identify to species. On some of
the Cumulative Panels in the water 9 months or more,
however, a few urchins of larger size were found and
could be assigned to one or the other of the two common
species of Strongylocentrotus. Small ophiuroids (Ophi-
othrix sp.) were also encountered on several of the Long
Term and Cumulative Panels.
Chordata (Tunicata):
Three species of tunicates were found on some of the
Long Term and Cumulative Panels, but invariably in
small numbers. Ascidia ceretodes and Styela truncata ap-
peared on panels, other than those in the intertidal rack,
that had been in the water 3 months or longer. The latter
were always small (5 mm or less) but specimens of Asci-
dia ceretodes on the 12 month panels were 50mm in
diameter. Aplidium solidum appeared on only a few
panels in the water 6 months or more. On an intertidal
panel in the water 12 months, specimens of Aplidium
solidum attained a diameter of 50 mm.
IV, ECOLOGICAL SUCCESSION
on LONG TERM PANELS
As noted earlier in the paper, during the year’s work
reported on here a rack of panels called Long Term
Panels was submerged one foot below low tide level. Some
THE VELIGER
Vol. 12; No. 2
of these panels (3-L-1, 3-L-2, etc.; see Figure 1) were
immersed on the first of each month and removed 3
months later. Another group of 4 panels exposed at 3
month intervals remained in the water for 6 months.
Using these panels it was hoped to determine something
regarding ecological succession. Specifically, these panels
were exposed to determine if the same or different popu-
lations of animals would collect during different 3 and 6
month periods of submersion during the year.
The 3 month panels developed at the end of their
period of submergence essentially the same populations
of animals regardless of the month when initially sub-
merged. Encrusting bryozoans dominated by Celleporaria
brunnea, but including Tubulipora pacifica and Crypto-
sula pallasiana practically covered each 3 month panel,
and a panel removed in January looked very much like
one removed in April or September. The colonies of these
bryozoans were smaller in size in the winter months, how-
ever, and this can be seen from the data for total biomass
(bottom line, ‘Table 2). Minimum biomass accumulated
during the November 1 to February 1 period (5.0 g per
panel side) and the maximum accumulated during the
May | to August | period (22.4 g per panel side). This
difference was due mainly to larger and thicker bryozoan
colonies in summer rather than more individual colonies.
ScHEER (1945) found that in the harbor at Newport,
California, a definite ecological succession occurred on
submerged panels involving a bacteria-diatom-bryozoan-
mussel sequence. In his studies a panel exposed in Decem-
ber went through the same sequence as one exposed in
March or April. He found that the time relations varied,
but the sequence did not. The panels exposed in this study
in Monterey Harbor were apparently not submerged long
enough to develop a final climax community, but the
evidence indicates that a true ecological succession such
as SCHEER observed is involved.
The 4 6-month panels exposed at different 6-month
periods during the year collected essentially the same
kinds of animals as the 3 month panels. Again, the en-
crusting bryozoans, especially Celleporaria brunnea, dom-
inated the community, but on a panel exposed from April
1 to October 1 and another exposed from July 1 to
January 1, the erect bryozoan Crisulipora occidentalis
formed clusters up to 4m high and contributed signifi-
cantly to the total biomass. The evidence indicates that
the community developed. by the end of 3 months does
not change significantly during an additional 3 months
exposure.
As can be seen from the data at the bottom of Table 2,
the total accumulated biomass on the 6 month panels
showed some strange variations. During the periods Oc-
tober 1 to April 1 and April 1 to October 1, the panels
Vole 12; No. 2
THE VELIGER
Page 191
collected a total dried biomass of 56.8 g and 61.3 g re-
spectively. Yet in the 6 month periods January 1 to July
1 and July 1 to January 1, the panels collected roughly
only half this biomass, namely 25.1 g and 29.7 g.
WOOD BORING ORGANISMS
During 1967-1968 the same two species of wood borers
were encountered as in the previous year. On the Short
Term Panels, Limnoria quadripunctata was found through-
out the year, but there were usually fewer than 10 gribbles
on any one panel and many of these were moving freely
over the panel surface or were just beginning to excavate
a burrow. On an intertidal panel removed after one
month on May 15, 1968, an adult and several young but
free gribbles were seen in a shallow burrow. The tiny
young had not yet begun to burrow. This is the first
instance in these studies where recently released young
Limnoria have been seen. At one time or another gribbles
were found on panels exposed at all depths, but they
preferred the intertidal panels or those submerged in the
shallow position. On the Long Term and Cumulative
Panels L. quadripunctata was found on every panel. On
those exposed for 3 months, up to 25 mature animals
could usually be found in burrows under the bryozoan
colonies, yet only the intertidal and shallow panels ex-
posed for 12 months showed extensive damage to the
wood. On some of these, however, the outer lamella of
the plywood was often riddled with Limnoria burrows.
The second wood borer, Bankia setacea, was found on
many of the Short Term Panels as tiny calcareous bodies
0.2 to 0.3mm in diameter with 2 siphonal apertures
separated by a calcareous bridge, but no siphons or
pallets were developed. These had barely penetrated the
wood surface. QuayLE (1959) has found that Bankia
attains this size and morphology within 4 days after
initial penetration of the wood. Never were more than 5
Bankia seen on any one panel, and the borer settled
mainly on shallow and deep panels, but on one occasion
on an intertidal panel. As can be seen from Table 1,
Bankia settled on Short Term panels in October, Decem-
ber, January, February, April and July. There was no
one period of maximum settling in 1967-1968. The range
of months over which settling occurred was broader than
has been found to be true in the open water of the bay
(Haperuiz, 1968b) where settling occurred from Decem-
ber through June.
In the Long Term and Cumulative Panels, Bankia was
also found, but rarely, and usually only 1 or 2 small
specimens per panel. Even panels exposed for 12 months
harbored few Bankia, yet the 1-inch redwood boards of
the shallow and deep racks were riddled with. burrows
of these shipworms at the end of 12 months, exposure.
This indicates that } inch douglas fir plywood is a poor
collector for Bankia. A project now in progress in Monte-
rey Harbor using 1 inch and 4 inch thick boards shows a
far greater abundance of Bankia than would be suspected
from past studies using plywood panels as collecting’ sur-
faces. Results of these studies on borers will be reported in
due time.
SUMMARY
(1) In continuation of a study initiated in Monterey
Harbor in 1966, a series of plywood panels were exposed
to the marine environment under Monterey Municipal
Wharf No. 2 at positions in the intertidal, one foot
below and 14 feet below low tide level. The period of
study in this report was from October 1, 1967, to Octo-
ber 1, 1968, with certain panels being observed through
December, 1968.
(2) Certain panels (termed Short Term Panels) were
exposed at each of the 3 positions for one month
periods throughout the year. These panels made it
possible to determine season of settlement of at least
the pioneering fouling and boring communities. Other
panels (termed Long Term and Cumulative panels)
remained exposed for from 3 to 12 months. These were
immersed at various times throughout the year and
made it possible to measure the rate of growth of many
organisms, to observe ecological succession and to de-
termine the nature of the dominant fouling community.
(3) A total of 66 species of animals settled on the Short
Term Panels during the year, while on the Long Term
and Cumulative Panels a total of 88 different species
were encountered.
(4) Encrusting and erect bryozoans were the dominant
fouling animals on all the panels. These included Celle-
poraria brunnea, Tubulipora pacifica, Cryptosula pal-
lasiana, and Crisulipora occidentalis.
(5) In contrast to the 1966-1967 year of study, few
acorn barnacles were recorded during this past year
and no period of mass settling was observed.
(6) As before, serpulid worms (Spirorbis spp. and Chi-
tinopoma occidentalis) were the most common annelids.
(7) Long Term Panels exposed for 3 and 6 months at dif
ferent periods throughout the year collected essentially
Page 192
THE VELIGER
Vol. 12; No. 2
the same population of animals regardless of the
month of initial exposure. The dominant forms were
encrusting bryozoans.
(8) The gribble Limnoria quadripunctata was commonly
seen on and burrowing into the plywood panels through-
out the year, but in small numbers. The shipworm
Bankia setacea was also recorded in small numbers in
many of the submerged panels, but there is evidence
that }-inch plywood is not a suitable collector of ship-
worms and does not give a true indication of the
number present.
LITERATURE CITED
Haper.iz, Eucene Ciinton
1968a. Marine fouling and boring organisms in Monterey Har-
bor. The Veliger 10 (4): 327-341; plt. 49; 3 text figs.
(1 April 1968)
1968b. Marine boring and fouling organisms in open water of
Monterey Bay, California. pp. 658-679 In: Biodeteriora-
tion of materials; microbiological and allied aspects. A. H.
Watters & J.S.Etprick, Eds. Elsevier Publ. Co., Barking,
England
MILLER, THomas LERoy
1966. Marine fouling organisms in Monterey Harbor, Cali-
fornia. Unpubl. Master's Thes. U.S. Naval Postgrad.
School, Monterey, Calif. (October 1966)
MomMsEN, DurwarD BELMONT
1966. A study of marine fouling in Monterey Harbor.
Unpub. Master’s Thes. U.S. Naval Postgrad. School, Monte-
rey, Calif. (May 1966)
Quay _e, Danie BraNcH
1959. The early development of Bankia setacea Tryon.
pp. 157-174 In: Marine boring and fouling organisms.
Washington Univ. Press, Seattle
ScHEER, BrapDLey Titus
1945. The development of marine fouling communities.
Biol. Bull. 89 (1): 103 - 121
Usnakoy, P V.
1955. Polychaeta of the far eastern seas of the U.S.S.R.
Zool. Inst. Acad. Sci. U.S. S.R., Moscow
Vol. 12; No. 2
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Page 193
Population Characteristics of Protothaca staminea (ConrAD)
from Mugu Lagoon, California
RONALD R. SCHMIDT
Department of Geology, University of California, Los Angeles, California 90024
AND
JOHN E. WARME
Department of Geology, Rice University, Houston, Texas 77001
(9 Text figures)
INTRODUCTION
STUDIES OF AGE STRUCTURES provide estimates of popula-
tion characteristics such as growth rate, average and
maximum longevity, and mortality rate. Protothaca sta-
minea (Conrab, 1837), the little-neck clam, is one of the
species of mollusks particularly useful for population
studies because it exhibits accentuated growth lines on the
surface of the shell. These can be interpreted as annual
rings, giving an age in years and a record of growth rate
for each shell.
The purpose of this study is to compare age and size
characteristics for a live population of Protothaca stami-
nea from Mugu Lagoon with the same properties for the
accumulating dead shells. Analysis of the dead shells pro-
vides information on mortality rate and the effect of
predatory gastropods on this species.
How population dynamics are reflected in the size-
frequency distributions of modern and fossil shell deposits
is a primary concern of paleoecology. Of the several
factors that contribute to the final size composition of an
in situ accumulation of shells, growth rate and mortality
rate are the most important. For our population of Proto-
thaca staminea in Mugu Lagoon, the size-frequency dis-
tribution of the dead shells is bimodal: a high peak of
juveniles and a smaller peak of adults. This is the conse-
quence of a high-to-low growth rate and, particularly, a
sigmoidal-shaped mortality rate not generally considered
in models of population dynamics.
SAMPLES rrom MUGU LAGOON
Location: The eastern arm of Mugu Lagoon (Figure 1),
just west of Point Mugu (34°30’N Latitude; 119°24’ W
Longitude), represents one of the few lagoonal environ-
ments along the southern California coast that has not
been seriously polluted nor altered by the activities of
man. The eastern arm is shallow, being nowhere more
than 3m deep at low tide. Hydrography is tidally con-
trolled and salinity essentially normal marine (approxi-
mately 34%). The general physical and biological aspects
of Mugu Lagoon have been treated elsewhere (WARME,
1966).
Dense populations of clams are present in most subtidal
and lower intertidal habitats in the lagoon. Our samples
were collected from a small channel between an island
and a mud flat (Figure 1). In order to get a representa-
tive sample of the shelled fauna, two parallel transects
were established normal to the length of the channel.
Five sample stations were picked along each transect at
intervals of about 10 m, and marked by semi-permanent
stakes. Profiles of the transects and the positions and
elevations of the samples are shown in Figure 2. Sampling
was completed on the afternoon of May 22, 1964, during
a period of low tide inside the lagoon.
Sampling Procedure:
A hollow cylinder with a cross-sectional area of {m’ and
a height of 75 cm, was used to facilitate sampling (Fig-
Page 194
THE VELIGER
Vol. 12; No. 2
Cc
Sampling Area Stations 1 to 10
100 meters 4, |Marsh
> Me i ie jee Se SS) L_» |
il te Sh N Barren
W en = a Ve? Zone
oe ay:
S [|Eel Grass
aia My
Mugu ) 4 (( The Eastern Region
ae
Va
v1 | ) =
Lagoon WIL | |
CoS \
rs ( . Wh
Ae a we ip
Pee S/,
Pacific Ocean
of Mugu Lagoon
600 meters
SSS SSS)
80 km
Santa ‘ :
_ Barbara (Ventura
, *~1 County
= TN
VENTURA \
t =" Los
N Mugu \_ -— Angeles
Lagoon ° 3
Figure 1
LOCATION MAP
Mugu Lagoon is situated on the coast of southern California
between Los Angeles and Santa Barbara (A). The sampling area
of Stations 1-10 is located in a channel between a small island
and a tidal flat in the eastern arm of Mugu Lagoon (B and C)
ure 3). This sampler can be pushed into loose sand and
mud by means of two handles, thus isolating equal areas
and known volumes of sediment at each sample location.
The sediment within the cylinder can be removed without
slumping of material from the sides.
Each sample was removed from the cylinder by shovel
and washed at the sample site to separate sand and mud
from living and dead shells. Washing was accomplished
with the aid of a screen with square openings, 3 mm on
a side, set in a wooden frame. The living and dead shells
thus collected were sorted, identified, and counted.
Three clam species dominated both the live and dead
assemblages in our samples: Protothaca staminea (Con-
RAD), Macoma nasuta (Conran, 1837), and Tagelus
californianus (Conrap, 1837). All were abundant enough
to be used for population studies (Figure 2), but only the
shell of PR staminea has readily discernible growth rings
Number of
live animals
Feet above
mean sea level
that can be used in estimating ages and growth rates.
Protothaca also has the advantage of a robust shell which
is not very susceptible to breakage during sampling.
SIZE anp AGE MEASUREMENTS
Sizes, defined as the greatest anterior to posterior length,
were measured with calipers to the nearest 0.1 mm.
Because many pelecypods add smaller increments of shell
during the winter months, their shells commonly show a
eo)
Number of
live animals
fo)
SB ®Ss
°
o}
Z
=
Om mM
5 é
Channel profile
Feet above
mean sea level
I 2 3 4 5
=. *'5 O° Meters =a
Cross-section of Traverse I
ma Protothaca staminea
aA Macoma nasuta
CJ Tagelus californianus
]
hannel profile
n
g
oD
S|
°
|
Histograms represent the three most abundant pelecypods of
Stations 1-10 in relation to Transects I and II. Elevations were
obtained by transit survey from a known benchmark. Low water
in the channel is just above mean sea level
Cross-section of Traverse II
Figure 2
crowding of growth lines or a “winter check.” These are
clearly visible on Protothaca staminea. “Annual rings”
were measured on each shell collected, and the measur-
ments indicating annual increments were recorded for
each year of growth. Most of the clams were less than 2
years old; thus data for the older and larger classes are
based on fewer measurements.
The “annual ring” method of age determination has
been discussed by WeyMouTH (1923) for Tivela stultorum
(Mawe, 1823); by Orron (1926) for Cardium edule
LinNAEuS, 1758; and by FRASER & SmitH (1928a) for
Paphia [= Protothaca] staminea (Conrap, 1837) and
(1928b) for Saxidomus giganteus (DESHAYES, 1839).
Vol. 12; No. 2
“Disturbance checks” may also be present; these can
sometimes be recognized and discounted, especially after
looking at many specimens from a given area. For some
species a plot of the class-frequency of all rings may help
to distinguish between disturbance and winter checks
(e. g., Craic & Hatiam, 1963, p. 738).
Figure 3
Sampling equipment used. The specially constructed cylinder, with
a 4m? cross-sectional area, was pushed 75cm into the soft sub-
stratum. Equal volumes (3/16 m3) of muddy sand were sieved at
each station
POPULATION CHARACTERISTICS
oF LIVE Protothaca
In the 10 samples collected in Mugu Lagoon, there were
180 live Protothaca staminea. Densities in individual
samples ranged from 1 to 31. The maximum values were
found in the deepest sample in each transect. These 2
samples(Figure 2, nos. 5 and 7) also included a greater
proportion of larger individuals.
Age and Size:
The distribution of ages within the total living popula-
tion is shown in Figure 4. There is a maximum number of
individuals which are up to one year old, and progressively
fewer individuals in the older year classes. The 7 year
time span represented in Figure 4 shows no clear “good”
or “bad” years in which reproduction or spatfall may
have been significantly greater or less than in other years.
The results of size-frequency measurements are pre-
sented in Figure 5. Truncation of the smallest classes
reflects the screen size openings used in collecting the
samples. The histogram is polymodal, the first peak re-
flecting the one-year or less class; the second peak is
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Page 195
60
he
aa 40
#30
I
°
o
Ee)
3
3
Z
erem sas 5. 6 078
Age in years
Figure 4
Age-frequency distribution of live Protothaca staminea,
Stations 1- 10, Mugu Lagoon
mostly the one and some two year olds; the third peak
is largely composed of two year olds with some three and
four year olds. The last peak represents individuals in
their fourth to seventh years. The position of the peaks
would be different if the samples had been collected at
other times during the year, being offset to the right as
growth proceeds. This would be evident particularly in
the faster growing younger classes. However, the overall
geometry of the curve should remain skewed as shown in
Figure 5.
Number of specimens
40 50
o 3
Length in mm
Figure 5
Size-frequency distribution of live Protothaca staminea,
Stations 1 - 10, Mugu Lagoon
Average Growth Rate:
The pattern of growth for Protothaca staminea is similar
to that known for other bivalves, with rapid early growth
and a steady decline in rate with age (HaLtam, 1967, p.
32). Figure 6 illustrates the average growth rate of P
staminea collected from the eastern arm of Mugu Lagoon.
With the exception of the first year, the growth rate of
P. staminea for this particular locality in Mugu Lagoon is
Page 196
Length in mm
Bite § © 7 8B OD
Age in years
Figure 6
Average growth rate of living Protothaca staminea,
Stations 1- 10, Mugu Lagoon
considerably lower than that measured by FRASER &
SmitH (1928a) for localities around Vancouver Island,
British Columbia.
Migration and Food:
On analysis of age, size, and rate of growth for our
samples, size differences appear to be related to age and
not to rate of growth. The average rate of growth is the
same throughout the channel. There is a proportionately
greater number of older (and larger) clams in the deeper
part of the channel. This may be accounted for by the
fact that Protothaca staminea can move laterally at the
surface of the substrate by a see-saw or rocking motion,
leaving a U- or V-shaped depression on the surface.
These trails can be several decimeters long, and once
recognized in the field a little-neck clam will invariably be
found at one end or the other. The adult Protothaca may
be moving in response to a more favorable food source.
The center of the channel has the greatest water exchange
and is always covered with water. It may therefore rep-
resent the location of maximum food supply and place of
maximum feeding time for the clams compared to the
higher, intertidal parts of the channel.
Fraser & SmitH (1928a) made extensive population
studies of live Protothaca staminea from the beaches of
Vancouver Island, British Columbia. They found that Pro-
tothaca populations which are protected from currents
and waves tended to exhibit a relatively high standing
crop of juvenile and young adult individuals. In more
exposed localities the populations contained relatively
fewer young individuals whose growth was slow in their
earlier years, but which grew better than average in their
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Vol 2s Now2
later years. Such exposed places may have a greater food
supply, but the young have trouble establishing themselves
owing to currents and waves. With growth the larger
clams are able to migrate to places with better food sup-
ply (FRASER & SmiTH, 1928a, pp. 265 - 266). A similar
migration may account for the distribution of larger
Protothaca in our samples.
Although no analyses have been made of the food of
Protothaca staminea in Mugu Lagoon, SmirH (1928, pp.
288 - 291) reports that this species is a suspension feeder,
collecting everything in the plankton small enough to be
ingested.
POPULATION CHARACTERISTICS
DERIVED From EMPTY SHELLS
Dead shells in our samples were about twice as numerous
as live individuals, reflecting a larger time of accumula-
tion compared to the time represented by the collected
standing crop. There was a good correspondence between
the number of left and right valves from these samples
(51% and 49%, respectively) even though the ligament
that holds Protothaca staminea valves together deterio-
rates when they die, and the valves separate if disturbed
a few weeks or months after death. Most of the discrepancy
between left and right valves occurred in the smaller
valves. These are more easily transported by currents and,
during sampling, are more likely to break and pass through
280
240 Bees
L. V.=334
WAAR. V.=327
ig Bored = 52
Valves
200 BS
Number of specimens
ao 4 oo 7
Age in years
Figure 7
Age-frequency distribution of the left and mght valves
(L. V. and R.V.) of dead Protothaca staminea,
Stations 1-10, Mugu Lagoon
Vol. 12; No. 2
the screen undetected. The amount of broken Protothaca
shells in our samples was minimal. Commonly it is diffi-
cult to ascertain if breakage is natural (by predation) or
whether it occurred during the sampling process.
Ideally, a dead clam would consist of left and right
matching valves (LV and RV in Figures 7 and 8). A
survey of the empty shells that were 2 years and older
100
80
'g 60
& YL. V.=334
H 40 AR. V.=327
2 Bored —
3 BB yeives 5?
20
10 20 30 40
Length in mm
Figure 8
Size-frequency distribution of the left and right valves
(L. V. and R.V.) of dead Protothaca staminea,
Stations 1- 10, Mugu Lagoon
indicates that most individuals are represented by corre-
sponding pairs. For our purposes, the number of dead
bivalves for each year or size class was derived from the
maximum frequency of either left or right valves.
Age and Size:
Figure 7 depicts the age distribution of mortality based
upon the annual rings on 661 empty valves collected in
Mugu Lagoon. Most deaths occurred during or at the
end of the first year of growth. In the older years, about
2 of the dead individuals collected have a winter check at
the margin of the shell. This indicates a higher mortality
during the winter months of decreased or non-growth, and
has been observed elsewhere for other temperate-climate
species (Craic & HALiam, 1963, p. 748).
The size-frequency distribution of dead shells is shown
in Figure 8. As was the case of the live clams collected,
truncation of the smallest classes reflects the size of open-
ings in the screen used as a collecting device. The first
and second peaks in Figure 8 correspond to mortality of
first and second year bivalves respectively. The paucity
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Page 197
of 2- to 5-year-old individuals accounts for the low num-
bers in the 20mm to 35 mm range (compare Figures 7
and 8). The peak centered around 40 mm represents a
relatively greater number of dead clams which were 5
years or older upon death.
Predation:
The carnivorous gastropods Forreria belcheri (Hinps,
1844) and Shaskyus festivus (Hinps, 1844) have been
observed preying upon Protothaca staminea and other
burrowing clams in and near the channel sampled in
Mugu Lagoon. About 15% of the dead P staminea
sampled had been drilled by boring gastropods (Figures
7 and 8). The predator seems to be somewhat selective
as to which valve it bores (34 LV and 18 RV).
On the average the predators appear to select shells
somewhat larger than the population mean: the mean size
of 52 bored valves was 17.3 mm, whereas the approximate
mean size of the remaining unbored individuals was 11.2.
This apparent selection could possibly be accounted for by
the chances that (1) the longer the bivalve is alive the
more likely that it will be preyed upon; (2) older clams
are more likely to have less resistance or be more attrac-
tive (ce. g., gonad areas are preferentially drilled in some
bivalves) ; and (3) a larger clam would be more likely to
be encountered by a wandering gastropod. If predation
were eliminated, the size classes of Figure 8 would be
shifted to the right (see Craic, 1967, p. 43).
POPULATION DYNAMICS
Interpretations of the dynamics of modern or fossil pop-
ulations derived from size-frequency distributions for shells
will depend largely on the assumptions adopted regarding
growth rate and mortality rate (e.g., Boucot, 1953;
Otson, 1957; FacersTroM, 1964; SHELDON, 1965). The
possible interactions between growth rate, mortality rate,
and the other less important factors, such as seasonal
recruitment of young and growth stoppages due to sea-
sonality or spawning periods, make many kinds of size-
frequency distributions possible in nature (Craic & OrR-
TEL, 1966).
Most invertebrate populations exhibit an average
growth-rate curve similar to Figure 6, with rapid growth
in youth and a decrease in rate with age. They also
commonly produce many more offspring than can possibly
survive to adulthood, and most of which die or are preyed
upon as larvae or juveniles. It is necessary that only a few
individuals reach sexual maturity for the population to be
maintained.
Page 198
Models of population dynamics usually assume morta-
lity rates that decrease, increase, or remain constant
with age (DEEvEy, 1947; Craic & OrrTEL, 1966; see
Figure 9). Assuming that dead shells have been neither
removed nor added at the site of deposition, a mortality
rate for Protothaca staminea can be calculated from the
age-frequency distribution of dead shells (Figure 7) and
presented as a log survivorship curve (Figure 9). Our
data suggest that the mortality rate for P staminea varies
log survivors
uw A BS A B.o ¥
Age in years
Figure 9
Log survivorship curve of Protothaca staminea, based on the dead
valves from Stations 1 - 10, Mugu Lagoon. Actual curve is compared
to types of mortality rate generally considered in population dynam-
ics: (a) increasing, (b) constant, and (c) decreasing
with age in a sigmoidal fashion. The risk of death is
high in the larvae and juveniles but is considerably
lowered upon sexual maturity. First spawn is not until the
end of the second or third year (FRASER & SMITH, 1928a;
Quayte, 1943). The mortality rate does not rise again
until individuals approach “old age,” forming what Com-
FORT (1957, p. 221) has described as a “plateau of adult
vigor.” In size-frequency distributions of accumulating
shells, this pattern of mortality would be reflected by a
bimodal distribution: a high peak of young individuals
and a smaller secondary peak of gerontic deaths (see
Figure 8).
Similar patterns of mortality have been described for
other mollusks (PELSENEER, 1934; ComFort, 1957).
FircH (1964) reports a population of Pismo clams,
Tivela stultorum, in which only 6 out of 200 live individ-
uals were less than 16 years old, and in which most empty
shells exceeded 5 inches (12.7 cm) in length, apparently
having succumbed to old age.
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Vol. 12; No. 2
CONCLUSIONS
Boucot (1953) described two theoretical size-frequency
distributions that may be of use in studying fossil popu-
lations. The first is right-skewed and represents a gradual
accumulation of shells of a stable population having high
juvenile mortality. Except for the secondary “old age”
peak exhibited by Protothaca staminea, the right-skewed
curve of Boucor and our Figure 8 are similar. Another
curve, approximating a “normal” or bell-shaped curve,
would theoretically result from current sorting or selective
destruction of empty shells by physical or diagenetic
processes (Boucot, op. cit.; see discussion by HALLAM,
1967). In relatively quiet-water locations, such as the
part of Mugu Lagoon where our samples were collected,
intense reworking of dead shells by physical forces would
not be expected (WarMeE, 1969), and the bell-shaped
size-frequency distribution was in fact not realized for
the empty shells.
In comparison, some samples of live Protothaca stami-
nea collected by FRASER & SMITH (1928a, pp. 258 - 260)
are right-skewed or left-skewed, and some show a nearly
normal distribution, depending upon environmental con-
ditions, such as wave exposure and tidal currents. In Mugu
Lagoon, deep-burrowing bivalves such as Tresus nuttalli
(Conrab, 1837) live in patches where most specimens are
of the same year class, giving a normal size-frequency
curve for each patch. A normal distribution can also be
approximated if there is a low mortality in the settled
young, as reported by Craic « Hattam (1963, p. 737)
for a population of Mytilus edulis Linnazus, 1758.
These examples suggest that size-frequency distribution
of live bivalve populations can vary greatly under a wide
range of ecological and environmental conditions. A nor-
mal distribution does not necessarily indicate a post-
mortem process working on the shells.
However, it is the mortality rate that has the ultimate
influence on age and size distributions of accumulating
shells that are potentially preservable as fossil populations.
Data concerning this vital statistic are rarely available
and, in the case of fossil populations, only attainable in-
directly. In our example, the size-frequency distribution
of empty Protothaca staminea shells in Mugu Lagoon in-
dicates that juvenile and gerontic deaths are most fre-
quent. The living standing crop, on the other hand,
contains a proportionately significant number of young
adults. It would appear that the death rate decreases upon
sexual maturity and does not rise until old age.
An interpretation of population dynamics depends upon
some means of determining age and growth rate, such as
Vol. 12; No. 2
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Page 199
growth-ring analysis. In addition, the empty shells yield
significant information concerning the distribution of
mortality that cannot be obtained from the live individ-
uals alone. An examination of age structures for both the
live individuals and the accumulating dead shells provides
an integral picture of the population dynamics of the
species under study.
ACKNOWLEDGMENTS
We express sincere gratitude to Dr. E. W. Fager, Scripps
Institution of Oceanography, University of California, San
Diego, for helpful criticism of the manuscript; however,
we are solely responsible for the content of this paper. We
wish to thank the U.S. Navy for access to Mugu Lagoon.
Margaret Rogers Schmidt drew the text figures accom-
panying this paper.
LITERATURE CITED
Bovucot, ARTHUR J.
1953. Life and death assemblages among fossils.
Journ. Sci. 251 (1): 25-40; 11 figs.
Comrort, ALEXANDER
1957. The duration of life in molluscs.
London 32 (6) : 219 - 241; 2 figs.
Craic, Gorpon Y.
Amer.
Proc. Malacol. Soc.
1967. — Size-frequency distributions of living and dead popula-
tions of pelecypods from Bimini, Bahamas. B. W.I.
Geol. 75 (1): 34-45; 12 figs.
Craic, Gorpon Y. « A. HaLLaM
1963. Size-frequency and growth-ring analyses of Mytilus edulis
and Cardium edule, and their palaeoecological significance.
Palaeontol. 6 (4): 731-750; 10 figs.
Craic, Gorpon Y. « GERHARD OERTEL
Journ.
1966. | Deterministic models of living and fossil populations of
animals. Quart. Journ. Geol. Soc. London 122: 315 - 355;
19 figs.
DeEvey, Epwarp S.
1947. Life tables for natural populations of animals.
Rev. Biol. 22 (4): 283-314; 9 figs.
Quart.
FAGERSTROM, J. A.
1964. Fossil communities in paleoecology: their recognition
and significance. Bull. Geol. Soc. Amer. 75 (12): 1197 to
1216; 5 figs.
Fitcu, JoHN Epcar
1965. A relatively unexploited population of Pismo clams,
Tivela stultorum (MaweE, 1823) (Veneridae). Proc. Mala-
col. Soc. London 36 (5): 309 - 312; plt. 13
Fraser, C. McLean & Gertrupe M. SMITH
1928a. Notes on the ecology of the little neck clam, Paphia
staminea CONRAD. Trans. Roy. Soc. Canada 22 (5): 249 to
269; 2 figs.; 5 plts.
1928b. Notes on the ecology of the butter clam, Saxidomus
gigantcus DESHAYEs. ‘lrans. Roy. Soc. Canada 22 (5):
271 - 286; 2 plts.
Hata, A.
1967. ‘The interpretation of size-frequency distributions in mol-
luscan death assemblages. Palaeontol. 10(1): 25-42;
11 figs.
Oxson, Everett C.
1957. Size-frequency distributions in samples of extinct or-
ganisms. Journ. Geol. 65 (3); 309 - 333; 17 figs.
Orton, Joun H,
1926. On the rate of growth of Cardium edule. Part. 1.
Experimental observations. Journ. Mar. Biol. Assoc. U. K.
14; 239-279; 12 figs.
PELSENEER, PAUL
1934. La durée de la vie et l’age de la maturité sexuelle chez
certains mollusques. Ann. Soc. Zool. Belg. 64: 93 - 104;
1 fig.
Quayte, DaniEL BRANCH
1943. | Sex, gonad development and seasonal gonad changes in
Paphia staminea Conran.
6 (2): 140-151; 5 figs.
SHELDON, RaymMonp W.
1965. Fossil communities with multi-modal size-frequency
distributions. Nature 206 (4991): 1336 - 1338; 6 figs.
SmitH, GERTRUDE M.
1928. Food material as a factor in growth rate of some
Pacific clams. Trans. Roy. Soc. Canada 22 (5): 287 - 291
WarME, JOHN E.
1966. — Paleoecological aspects of the Recent ecology of Mugu
Lagoon, California. Ph. D. dissertation, Univ. Calif. at
Los Angeles: 380 pp.; 41 figs.; 5 plts.
1969. Live and dead molluscs in a coastal lagoon.
Paleontol. 43 (1): 141-150; 2 figs.
WEYMOUTH, FRANK W.
1923. The life-history and growth of the Pismo clam (Tivela
stultorum Mawe). Fish. Bull., Calif. State Fish & Game
Comm. 7: 120 pp.; 15 figs.
Journ. Fish. Res. Brd. Canada
Journ.
Page 200
THE VELIGER
Vol. 12; No. 2
Mimicry of the Gastropod Mitrella carinata
by the Amphipod Pleustes platypa
JULES M. CRANE, Jr.
Cerritos College, Norwalk, California 90650
(Plate 36)
THE PLEUSTID AMPHIPOD, Pleustes platypa BARNARD &
Given, 1960, was described on the basis of 3 preserved
specimens collected off Palos Verdes, Los Angeles County,
California (BARNARD & GivEN, 1960, pp. 41 - 42; fig. 1).
Because preserved specimens rapidly lose their color and
have their appendages in unnatural positions, it was not
apparent that in life these amphipods look remarkably
like the snail Mitrella carinata (Hinps, 1844). In May,
1966 a single living specimen of P. platypa was discovered
on the kelp Macrocystis pyrifera by the Cerritos College
Marine Biology class near Point Loma, San Diego, Cali-
fornia. Collected as Mitrella carinata, the amphipod char-
acteristic was not observed until much later when the
collection was examined in the laboratory. Because of the
uncertainty about the maintainance of this specimen alive,
it was observed briefly and then preserved.
When it became apparent that this was the only known
arthropod molluscan-mimic, efforts were made to obtain
more specimens for observation and photography. Finally,
on September 17, 1968, 5 living specimens were collected
from a Macrocystis holdfast floating at a depth of 15
feet off Point Dume, Los Angeles County, California.
These were photographed the same day using randomly
collected specimens of Mitrella carinata for comparison
(Plate 36).
Pleustes platypa tucks its pleon up under the pereon
where it is shielded from view by the enlarged and flat-
tened protopodites of the 6", 7", and 8" pereonal (tho-
racic) appendages and the enlarged coxal (lateral) plates
of the pereon. The 2 pairs of gnathopods are similarly
hidden by the coxal plates. The amphipod rests at about
a 30° angle, supported by the 6", 7", and 8" pereopods,
which also serve as locomotor structures in this position.
These appendages permit the amphipod to crawl slowly or
launch rapidly into darting, short-distance swimming
activity.
The amphipod coloration is highly variable, as is the
coloration in Mitrella. Both specimens shown in Plate 36
have similar color patterns. This particular amphipod
had a dark grey cephalon, a yellow band around the
anterior half of the pereon, and graded from light brown
to dark brown posteriorly. The pereopods (walking legs)
often display light and dark grey or brown bands which
mimic the mottling found in the foot, siphon, and eye
stalks of Mitrella. These are contrasted with the darker
coloration of the protopodites.
Judging by skin divers’ reports and recorded captures,
Pleustes platypa appears to be rare but this is probably
because of the difficulty in distinguishing it from the
extremely abundant Mitrella carinata. All specimens have
been found in association with Macrocystis. The presently
known range of P platypa is herein reported as Point
Loma, San Diego, California to Gaviota Beach, Santa
Barbara County, California. A single observation of an-
other as yet unidentified species of amphipod mimicking
a different form of Mitrella carinata found in pholad
holes in cobble off Point Loma, California, suggests that
the whole range of polymorphism in Mitrella may be
reflected in mitrelliform amphipods.
ACKNOWLEDGMENTS
I wish to thank Dr. Rimmon Fay, president, Pacific Bio-
marine Company, for providing the live specimens; Dr.
J. L. Bamard, U.S. National Museum, for the identifi-
cation of the initial specimen; and Robert L. Eberhardt,
Lockheed Aircraft Corporation, for cooperation in the
use of the RV Sea Quest. The photograph of the living
animals was taken by George Reeves.
LITERATURE CITED
BarNARD, J. LAURENS & Ropert R. GIVEN
1960. Common pleustid amphipods of southern California,
with a projected revision of the family. Pacif. Naturalist
1 (17): 41-42; 1 text fig. (15 April 1960)
Tue VELIcER, Vol. 12, No. 2 [CRANE] Plate 36
Mitrella carinata (left) showing mimicry by the amphipod Pleustes platypa (right)
i
Vol. 12; No. 2
THE VELIGER
Page 201
Cypraeidae of the Red Sea at Massawa, Ethiopia,
with a Zoogeographical Analysis
based on the Scuitpers’ Regional Lists
BY
T. C. FOIN
Institute of Ecology, Environmental Systems Group, University of California, Davis, Davis, California 95616
AND
L. P RUEBUSH
Box 67 USA STRATCOM Fac., Asmara, APO New York 09843
(3 Text figures; 1 Table)
BECAUSE OF THE INTERESTS Of shell collectors, the genus
Cypraea has been the subject of an expansive literature,
including many lists of species from various faunal prov-
inces, To our knowledge no list of Cypraea from the Red
Sea since the compilation of ScHILDER & ScHILDER (1938-
1939) has been published, however, despite the interesting
species that have resulted from the isolation of the Red
Sea from the remainder of the Indian Ocean basin. The
SCHILDER & SCHILDER records were based on literature
records, and many of their recorded species require con-
firmation. This paper represents several years of collecting
by the junior author in the southern Red Sea, and is a
compilation of the species found in the area with a com-
parison to other Indian Ocean species.
The Massawa Area
Massawa, Ethiopia, is located at approximately 39°29’ E
Latitude and 15°39’ N Longitude on the western shore of
the Red Sea. The Dahlak Archipelago completely shields
Massawa from the open sea to the east, but Massawa is
an important port because of the two channels that pene-
trate the Dahlak Archipelago from the north and the
south. The remainder of the area is shallow and generally
does not exceed 5 fathoms in depth. Cypraea spp. are
found in many habitats, ranging from mudflats to coral
reefs; some species are accessible intertidally, but most are
obtainable only by diving. Most collections were made in
this manner. Collections were made in all seasons; the
only notable changes in species abundance occur in the
winter months, when it is assumed that many individuals
either hide and are inactive, or migrate to deeper water.
tomical studies in this paper are limited to color obser-
All measurements were taken with vernier calipers; ana-
vations and gross detail of external structures. A map of
the collecting area is presented in Figure 1.
Cypraea of the Southern Red Sea
Cypraea annulus LinNAEuS, 1758
This species is very common in the low intertidal and
shallow subtidal zones, restricted to areas of old coral
and rock rubble possessing strong currents. Shell form is
in agreement with that given by Orr (1959) for Cypraea
annulus from similar habitats off Zanzibar.
Cypraea arabica LinNAEus, 1758
(Cypraea grayana ScHILDER, 1930)
This species is widespread over both the habitat in which
Cypraea annulus is found and in living coral. Specimens
appear to be more abundant in the former habitat. All
specimens found have been small for the species, averaging
about 30 mm. Cypraea arabica is one of the most variable
Page 202
39° 30" 40’
Daret Island 4 4. \,
Dehel Island
50
Ras el Garara
C Massawa
Sheikh Said Island
a7 Ras Gedem
oles
Gedem Mountain 34
30
Eritrea
Gulf of Zula
15°90’
THE VELIGER
Dissei Island
p Dur Ghella Island Q ce .
Dur Gaarn Island
rts)
Ras Arb
g,
» Nocra Island
Ghubbet
Mus Nefit
a
Enteara Island
o
Madote Island
Buri Peninsula ©." =
a
fo) 10
Figure 1
Map of the Massawa, Ethiopia area, including a part of the
Dahlak Archipelago
Adapted from a U.S. Army Map Service chart made in 1936
species found in the Massawa area; specimens may be
found that resemble C. grayana, C. eglantina Ductos,
1833, and C. arabica. The diagnosis given by SCHILDER &
SCHILDER in their prodrome for C. grayana emphasizes a
subpyriform, often humped shape, reticulate or dilacerate
dorsal pattern, and a spire blotch. The ScuitpErs and
CrernoHorsky (1964) diagnose C. eglantina as having
a cylindrical shape, unmargined sides, a spire blotch, and
a dorsal pattern of longitudinal lines interrupted by pale
lacunae. Cypraea arabica is said to differ from these 2
species by the lack of a spire blotch, its hieroglyphic dorsal
pattern, ovate shape, and the heavily margined sides. Our
analysis of the Massawa “arabica” (Figure 2) of the
above characters is based on a classification of each shell
Vol: 12;3Now2
20 miles
Vol. 12; No. 2
with respect to the base, dorsum, shape, and spire blotch.
In the logical tree only the combinations found of the
36 possible are presented. Only one of the sample (N =
19) examined was a C. grayana and 3 were C. arabica;
Spire
Blotch Base Shape Pattern No. Specimens
rp margined —— subpyriform — reticulate I
present
Lrounded — subpyriform — reticulate 1 (= grayana)
hieroglyphic 3 (= arabica)
ovate — longitudinal 3
reticulate I
eon hieroglyphic 2
Schley ae longitudinal 4
absent — pa
cylindrical — hieroglyphic 2
rounded |
subpyriform — hieroglyphic
Nn
Figure 2
Distribution of shell characters of a sample of
“Cypraea arabica” from Massawa
of the remainder, all but one most closely resembled C.
arabica. We have, therefore, provisionally called the Mas-
sawa specimens C. arabica instead of C. grayana. This
analysis also suggests that a reexamination of the group of
species in the “arabica” complex may be warranted.
Cypraea camelopardalis Perry, 1811
Cypraea camelopardalis is relatively common throughout
the region on and under coral reefs. This species is the
most variable local species in size and pattern; the smallest
adults range down to 35mm, while the largest exceed
70mm. The dorsum is frequently heavily scarred. Two
adult patterns may be distinguished: the normal pattern,
with white spots scattered over the dorsum; and individ-
uals ranging from a few white spots to none at all. These
differences are assumed to be both genetic and environ-
mental, but since all combinations have been found to-
gether, these patterns are thought to bear no taxonomic
significance. The color of the external anatomy of C.
camelopardalis is the same as in C. vitellus LINNAEUS,
1758, as described by CeRNoHorSKY (1964); this is
further evidence of the supposed close relationship of the
two species.
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Page 203
Cypraea carneola LINNAEUS, 1758
Cypraea carneola, like C. annulus, is restricted to areas of
rock and coral rubble swept by strong tides and currents,
but may be locally abundant in these places. All species
found in these areas have been characterized by small
size; C’. carneola is no exception. Specimens average about
25 mm in size. The dorsum normally is heavily scarred.
Conchologically, anatomically, and ecologically, speci-
mens found in the Massawa area closely resemble typical
C. carneola from the Indo-Pacific.
Cypraea caurica LINNAEUS, 1758
C'ypraea caurica may be locally common, although like C.
carneola, it is restricted to specific habitats. In the Massa-
wa area the species is usually found under isolated coral
heads, either living or dead. The species is quite variable
in size, in labial denticles, and in the presence or absence
of a dorsal blotch, but all specimens are easily recognized
as C’. caurica. The nominal subspecies C’. c. quinquefas-
ciata Born, 1778, inhabits the Red Sea. We have not
evaluated the validity of this name.
Cypraca clandestina LINNAEUS, 1767
A single living specimen of this species has been found
under a rock off Dissci Island, SE of Massawa near the
Gulf of Zula. Beach specimens are similarly very rare;
we infer that either this species does not normally occur
in the Massawa area or is very rare there.
Cypraea erosa nebrites MELvILL, 1888
In the Massawa area individuals of Cypraea erosa neb-
rites are mostly restricted to the islands, largely in areas
of isolated living coral and rock rubble, although a few
have been found on the mainland at Ras El Garara.
Cypraea e. nebrites is easily recognized: the marginal blot-
ches are always prominent, do not extend unto the base,
the teeth are distant and streaked with red, and the shell
is more rounded and compact than Indo-Pacific C’. erosa
Linnaeus, 1758. The animal also differs in color from
Fijian examples: there is a predominance of green in the
mantle lobes and siphon. Cypraea e. nebrites is relatively
common, but Massawa examples are smaller than those
from more northerly localities.
Cypraea exusta SOwERBY, 1832
Cypraea exusta is restricted to the NW Indian Ocean
and is very rare in the Red Sea. Only 5 specimens have
been found, all together, under a coral head in the south
Page 204
harbor of Massawa. Specimens are easily distinguished
from C. talpa Linnaeus by the less expanded base, the
more rotund shape, the extremely dark coloration, and
the very fine and numerous columellar teeth. It is possible
that C. exusta is a rare subspecies of C. talpa; at least, they
are sibling species.
Cypraea felina GMEun, 1791
Cypraea felina is one of the more common species in the
Massawa area; it may be found in rocky habitats, usually
in association with C, annulus, C. arabica, C. carneola,
and C. caurica. This species is not very variable in size or
shape. Length-breadth-height measurements comparisons
with C, f. fabula Krener, 1843, from the Arabian Gulf
and C. f. felina of East Africa indicate that the Red Sea
subspecies is probably the latter.
Cypraea gracilis notata Giti, 1858
This species is common in the same habitats as Cypraea
felina. It occurs in two distinct shapes, one more cylind-
rical, the other pyriform. These forms may be found to-
gether, and no ecological significance has been attached
to this difference. The colors of the living animal feature
a white mantle, orange foot and head, and black eyes.
Average lengths of the Massawa individuals range from
13 to 16 mm.
Cypraca isabella LinNAEuS, 1758
Only a single example of Cypraea isabella has been found
living in the Massawa area (South Harbor). This species
does not normally appear to occur in the area.
Cypraea lentiginosa Gray, 1825
Cypraea lentiginosa is occasionally in the Dahlak Archi-
pelago but very rarely along the mainland. One was found
in the South Harbor. It is apparently a recent immigrant
to the Red Sea; ScHILDER & ScHILDER (1938-1939) list
its center of distribution as the waters between the Gulf of
Oman and the Indian subcontinent.
Cypraea lynx Linnagus, 1758
This species is occasionally found in the offshore islands
but rarely along the mainland. It occurs in the same hab-
itats in which Cypraea camelopardalis and C. pantherina
Licutroot, 1786, are found. The species is said to be
represented in the Red Sea by the subspecies C. 1. williamsi
Me vii, 1888. The ScuitpeErs give a diagnosis for C.l.
williamsi; they cite a short anterior terminal ridge, blunt
posterior extremities, rounded sides, obsolete basal carinae,
wide aperture, and a pink base. Base color cannot be
THE VELIGER
Vol. 12; No. 2
used as a subspecies criterion in this case, as C. lynx with
pink bases may be found haphazardly over the Pacific and
Indian Ocean basins, in all the other nominal subspecies.
Individuals of C’. lynx from various localities in the Indo-
Pacific also show the characteristics of C. J. williamsi;
we feel that C. /. willamsi is not a valid subspecies, but
this conclusion needs confirmation.
Cypraea pantherina Licutroot, 1786
Cypraea pantherina is one of the most common and con-
spicuous cowries of the Red Sea. It is widespread over the
region but is restricted to areas of coral development. This
species exhibits considerable variation in dorsum colora-
tion, from nearly pure white with few spots, to nearly all
black. There are also stages of development of a goiden
color form, which at its most extreme stage appears as a
solid golden brown (funebralis SuLuorti, 1924). In many
specimens the golden nacre is succeeded by a pattern of
black spots over white; the golden color persists as the
dorsal line. If this is also true for the darkest specimens,
the color may persist for less than the lifespan of an
individual. A large series of C. pantherina demonstrates
the intergradation in color that occurs; under these cir-
cumstances the name funebralis is no more than the name
of a specific color form, one which may last for less than
the adult life of an individual. Accordingly, it is probably
inappropriate that even the informal taxon forma should
be used for funebralis, and the name should be dropped.
Cypraea pulchra Gray, 1824
This species is found in coral in the Massawa area. The
individuals encountered are easily recognizable, as there
is almost no variation shown by the species, other than
in size.
Cypraea staphylaea LinnagEus, 1758
A few beach specimens of this Indo-Pacific species have
been found, but no living examples are known from
Massawa.
Cypraea stolida erythraeensis SowERBY, 1837
Cypraea stolida erythraeensis is restricted to coral reefs on
the offshore islands, although it occasionally occurs on the
mainland. This subspecies is considerably smaller than the
other subspecies of C. stolida; it averages only about 18mm
in length. Specimens are characterized by a plain white
base with teeth produced and a large, often isolated, dor-
sal blotch. Cypraea s. erythraeensis appears to be quite
invariable except for the form and dimensions of the
dorsal blotch. Tooth color is also said to be variable (R. H.
r
Vol. 12; No. 2
Summers, personal communication). The ScHILDERs ac-
cord specific status to C. s. erythraeensis.
Cypraea turdus LAMaRcK, 1810
Cypraea turdus is frequently found in company with
other species inhabiting areas of isolated rock and coral
and possessing strong tidal surges and currents. There is
considerable size variation, although all specimens seen
have been quite broad. An occasional specimen will be
nearly all white as the result of very extensive lateral
callouses. This species appears to be the most widespread
and common of the Massawa Cypraeidae.
The ScuiLpers’ list includes several species that have
not been found in the Massawa area. These are C'ypraea
cicercula LinNAEUS, 1758, C. macandrewi SOWERBY,
1870, C. nucleus Linnagus, 1758, C. helvola LINNAEUS,
1758, C. teuleri CAZENAVETTE, 1846, C. talpa LINNAE-
us, 1758, C. mappa LinnaEus, 1758, and C. mauritiana
LinnaEus, 1758. Of these, the ScHi~pERs expressed
doubt about validity of the C. talpa and C. mauri-
tiana records; because these are large and normally
common species, we are inclined to agree. We also do
not belicve that C’. moncta occurs in the Massawa area.
Some recent collections of C’. nucleus, C. macandrewi, C.
cicercula, and C. gangranosa have been reported from the
Red Sea (R. H. Summers, personal communication). We
infer that all these species are rarely collected in the
Massawa area. We have no information concerning C.
mappa, C. helvola, or C. teulerei, although we have
received one unconfirmed report on the former species.
Zoogeographical Implications
of the Red Sea Cypraea
The British Navy Intelligence Division (1946) has sum-
marized the geological history of the Red Sea basin. The
Red Sea is a shallow, warm-water, marginal sea of the
Indian Ocean formed by the flooding of part of the East
African rift valley around the Pliocene. Shortly thereafter,
the Red Sea and the Gulf of Aden were connected to the
Indian Ocean, and the main northward flow through the
narrow strait (Bab-el-Mandeb) that marks the mouth
of the Red Sea was established. One would expect that
the recent origin and isolation of the Red Sea would have
marked effects on the cypraeid fauna.
Eighteen species are listed from Massawa in this paper,
of which 5 are endemic to the Erythraean region. When
the regional lists of ScHILDER & SCHILDER (1938-1939)
were analyzed and doubtful taxa removed, the Erythrae-
an province clearly had a greater proportion of endemic
THE VELIGER
Page 205
species than any other province of the Indian Ocean
(Table 1). We removed doubtful records and lumped all
Table 1
Reduced number of species and proportion of endemic species
in regions of the Indian Ocean
Number Number
of species endemic
Northern Red Sea 19 8
Central Red Sea 24 9
Southern Red Sea 37 10
Gulf of Aden 42 10
Chagos Archipelago 43 1
Mauritius 49 1
St. Francis Bay to East London 27 6
Zanzibar 40 0
“sympatric subspecies” into a single species for these an-
alyses. Our records generally agree with the ScHILDERs’
list, and are evidence of the isolation of the Erythraean
region from the remainder of the Indian Ocean. We feel,
therefore, that the ScuILpeErs’ lists are sufficient for zoo-
geographical analysis. The cypraeid fauna is also depau-
perate in comparison to the remainder of the Indian
Ocean, but the total number of C'ypraea species is similar
to that of the South African region. Isolation has pro-
duced several nominal subspecies — nebrites, erythrae-
ensis, and notata — and 3 species — camelopardalis,
exusta, and pantherina. These 3 species are easily derived
as sibling species from 3 common Indo-Pacific ones. These
are Cypraea vitellus, C. talpa, and C. tigris, respectively.
When the reduced number of species in the SCHILDERs’
lists for the Erythraean region are plotted against distance
of the midpoint of the length of the eastern Erythraean
region (Gulf of Aden), Figure 3 results. When plotted as
a semilog or log-log function, the points more nearly fall
in a straight line, but a distinct inflection remains in the
curve. Two explanations are possible: either there are
more species in the Red Sea than can be accounted for
as a simple function of distance, or the Gulf of Aden is
depauperate. The latter explanation is unlikely as the
reduced number of species reported (43) compares with
the main basin of the Indian Ocean (Table 1). The
first report of Cypraea clandestina, C. gangranosa, and
C. lentiginosa from Massawa, and the increasing propor-
tion of regionally endemic species in a northward direc-
tion through the Red Sea, are evidence that species char-
acteristic of other Indian Ocean provinces are still enter-
ing the Red Sea, and that given sufficient time the number
Page 206
50
40
30
Number of Species
20
500 1000 1500 2000
Distance from Midrange of Gulf of Aden
Figure 3
Decreases in number of species as a function of distance
from the NW Indian Ocean to the Gulf of Suez,
from the ScHILDER & SCHILDER lists
of species in the Red Sea should equilibrate with the rest
of the Indian Ocean.
We interpret the inflection in the curve of Figure 3,
and the number of new species found at Massawa, to
mean that Bab-el-Mandeb is not an effective barrier to
species immigrating from the Gulf of Aden into the Red
Sea, at least at present. The presence of at least 6 sibling
species and subspecies implies that the Red Sea was
either formerly more isolated or that immigration of a
particular new species into the Red Sea has been sporadic
and infrequent. We cannot now differentiate between
these.
THE VELIGER
Vol. 12; No. 2
SUMMARY
(1) Eighteen species have been found in the Massawa,
Ethiopia area of the 27 reported by ScHILDER & ScHIL-
DER (1938-1939) list. Four of the 9 species remaining
to be collected are probably present but rare, and 3 are
probably erroneous reports.
(2) The Massawa Cypraeidae contain a high percentage
of regionally endemic forms (8 of 18), but 6 of these
are easily derived from widespread Indo-Pacific species.
(3) The number of endemic forms and the number of
Cypraea species indicate that the Red Sea was at least
at one time highly isolated from the remainder of the
Indian Ocean.
(4) Analyses of several nominal subspecies from the Mas-
sawa area and of the ScHILDER & SCHILDER lists suggest
that gene flow within the Cypraeidae may be wide-
spread.
LITERATURE CITED
CERNOHORSKY, WALTER OLIVER
1964. The Cypraeidae of Fiji (Mollusca:Gastropoda).
The Veliger 6 (4): 177-201; plts. 21-26; 1 Text fig.; 1 map
(1 April 1964)
NavaL INTELLIGENCE Division, Royat Navy, GREAT BRITAIN
1946. | Western Arabia and the Red Sea. Geograph. Hand-
book Ser. 527
Orr, VIRGINIA
1959. A bionomic shell study of Monetaria annulus (Gastro-
poda : Cypraeidae) from Zanzibar. Notul. Natur. 313
ScHILDER, FRANZ ALFRED
1962. Hybrids between Cypraea tigris Linnarus, 1758 and
Cypraea pantherina SoLANDER, 1786. The Veliger 5 (2):
83 - 87, 1 diagram. (1 Oct. 1962)
SCHILDER, FRANZ ALFRED, & MARIA SCHILDER
1938 - 1939. Prodrome of a monograph on living Cypraeidae.
Proc. Malacol. Soc. London, 23 (3): 119-180; (1939) 23
(4): 181 - 231; 1 text fig.; 9 maps.
(15 November 1938 and 15 March, 1939)
Vol. 12; No. 2
THE VELIGER
Page 207
The Distribution and Ecology of Sub-Littoral Species of
Macoma (Bivalvia) off Moresby Island and in Satellite Channel,
near Victoria, British Columbia
R. M. DUNNILL
AND
D. V. ELLIS
Department of Biology, University of Victoria, Victoria, British Columbia, Canada
(9 Text figures; 3 Tables)
INTRODUCTION
THE GENUS Macoma Leacu, 1819 (Bivalvia) is widely
distributed in northern seas and is also known to occur
south of the equator in East Africa (THorson, 1957, p.
106). It has undergone wide speciation, particularly in
the eastern North Pacific and DUNNILL & Etuts (1969)
have confirmed 13 species presently occurring in the shal-
low coastal waters of southern British Columbia. Previous
ecological studies (and incidental mention) of Macoma
in the eastern North Pacific have in general been con-
fined to the littoral species MM. inquinata (DESHAYES,
1854), M. nasuta (Conran, 1837), M. secta (Conran,
1837), (amongst others, SHELFORD & Tow er, 1925;
SHELFORD et al., 1935; MacGrnitic, 1935; AppicotTT,
1952). The dearth of detailed ecological studies on sub-
littoral species of Macoma probably stems from earlier
confusion over the taxonomy of the genus.
This paper discusses the distribution of 8 species of
Macoma occurring sub-littorally in an area of predomi-
nantly sandy sediments off the north end of Moresby
Island, near Victoria, British Columbia, and examines the
effects of some environmental factors.
Data from a previous benthic survey in adjacent Sat-
ellite Channel by Etxis (1967 and in press) are included
since this extends the range of sediment types covered.
' Based on part of a thesis submitted by R. M. Dunnill in partial
fulfillment of the requirements for the degree of M. Sc. at the
University of Victoria
ACKNOWLEDGMENTS
We wish to thank F Bernard, J. Dobrocky, R. Herlinveaux,
D. B. Quayle, R. G. B. Reid, and D. Thorndick for their
assistance in various aspects of this study.
Federal Government vessels were made available for
the surveys through the Canadian Committee on Oceano-
graphy.
The investigations were financed by the National Mu-
seum of Canada, the National Research Council of Cana-
da, the Defence Research Board and the University of
Victoria.
DESCRIPTION or tHE STUDY AREA
PHYSIOGRAPHY
Satellite Channel, on the south-western fringe of the Strait
of Georgia, southern British Columbia (Figures 1 and 2)
is roughly 13 nautical miles long, 1.5 to 2.0 nautical
miles wide and has a mean depth of 73m, although in
several places, narrow elongated zones occur with depths
greater than 90m. Oceanic water has access to Satellite
Channel from Swanson Channel and Haro Strait over a
sill at a depth of 66m.
Moresby Island lies at the east end of Satellite Channel
at its junction with Swanson Channel.
Page 208
THE VELIGER
™
Gulf Islands
~ WY
Gyn
Riya
Satellite Channel
San Juany
ee Op
Figure 1
@ Hydrographic Station \
e Anchor Dredge
4 Van Veen _
Time Series
0 1 2
Nautical Miles s
123°30’W
Figure 2
Satellite Channel and Moresby Island showing station localities
a amas
123°20°
Vol. 12; No. 2
Vol. 12; No. 2
HYDROGRAPHY
Tuxtty & Dopimeap (1957) and WatpicHuk (1957)
discuss the oceanography of the Strait of Georgia.
According to WALDICHUK (op. cit.), there are basically
3 water masses in the Strait of Georgia-San Juan system:
1. the brackish surface water from run-off in the Strait
of Georgia (mainly from the Fraser River) ;
2. the deep water of oceanic origin in Juan de Fuca
Strait; and
3. a mixture of (1) and (2) which forms at the sills
between the straits.
Tidal currents in the southern region of Georgia Strait
flow rapidly and turbulently in narrow passages between
mazes of islands (TuLty « Dopimeap, 1957). Intensive
March 31 1959 June 8 1959 November 16 1959
Station B B E
Temperature °C
pent 9 iowa a2 8 10
25
50 |
|
75 |
Salinity %
026 28 30 29 31 27
Depth in meters
w
=)
Dissolved Oxygen (m1/1!)
4 6 8
50
75
Figure 3
Water properties in Satellite Channel
THE VELIGER
Page 209
tidal action, aided by wind and waves, particularly in
winter, mixes the brackish surface layer with underlying
more saline water to near homogeneity. Temperature and
salinity stratification is minimal in the channels of the
southern Strait.
In Satellite Channel, in the northwestern portion of the
southern region, current velocities are lower, there is less
mixing and more stratification develops. Seasonal changes
in properties of the upper 75 - 80 m of the water column
in Satellite Channel are shown in Figure 3, based on data
from HERLINVEAUX et al. (1960) and Pacific Oceano-
graphic Group (1959a and 1959b).
A pronounced thermocline, established in summer, is
destroyed by increased mixing in winter. The deeper water
averages about one degree Centigrade lower in winter
than in summer.
Salinity changes mainly reflect the seasonal cycles of
run-off of the Cowichan River (HERLINVEAUX, 1962). A
prominent halocline develops in the upper 10 m in winter
and spring, coinciding with maximal annual precipitation
and run-off in the area at sea level. The surface lower
salinity layer is much less in evidence in summer, when
discharge from the Cowichan River is low. However, a
less pronounced halocline persists, possibly indicating the
presence of Fraser River outflow, which is maximal in
summer.
From 15-85 m, temperature, salinity, and dissolved
oxygen values show little seasonal or vertical variation.
Temperatures range from 7.87 -9.70° C, salinities from
29.25 - 31.46%, and dissolved oxygen concentrations from
4.37-6.13 ml/l. Ranges in these properties seem too
small for them to be important in limiting distribution of
sub-littoral Macoma species in Satellite Channel at
depths investigated. Water off the north end of Moresby
Island, closer to regions of intense mixing than the hydro-
graphic stations in Satellite Channel, probably is less
stratified and has smaller ranges in temperature, salinity,
and dissolved oxygen concentrations. Sediment tempera-
tures measured off Moresby Island in the summer of 1967
(Table 1) had a range of less than 1° C at stations
between 26.5 and 84.0 m.
GEOLOGY
Sediments in Satellite Channel and their Origins
There is a marked progression in the Satellite Channel
trough from very fine grained sediments at the western
end near the mouth of the Cowichan River to coarser
grained sediments in the east (Figure 4). The mean silt-
clay fraction was 95.4% in Cowichan Bay and 82.2% at
the mouth of the bay. Further east, the silt-clay fraction
had decreased to 40.3% at Satellite Channel Center and
Page 210
%MSilt-clay
O- 25
25- 50
50- 75
75 - 100
Shell gravel #48
0 il
| Nautical Miles
123°30’W
THE VELIGER
Vol. 12; No. 2
Figure 4
Generalized distribution of sediments in Satellite Channel
to 25.9% at Satellite Channel East. Excluding stations on
“shell-gravels”, the mean of the 13 stations (26 samples)
on the finer sediments in the grid at the north end of
Moresby Island was 31.3% silt-clay.
There were localized exceptions to this main trend.
Bosun Bank on the west side of Satellite Channel, near its
northwestern end, was noticeably sandy (15.3% silt-clay),
while Fulford Harbour near the east end had fine grained
sediments (69.2% silt-clay).
“Shell-gravels” occurred at 4 stations at the mouth of
Moresby Passage (Figure 2), where currents reach 2 - 3
knots (Canadian Hydrographic Service, chart 3452).
The fine grained sediments at the west end of Satellite
Channel, in the vicinity of Cowichan Bay, probably owe
their origin to material carried down in suspension by the
Cowichan River. It is improbable that the coarser grained
fractions of the sediments in the channel trough originate
from material brought down by the Cowichan River since
these would, in that case, be deposited before the fine
grained material in Cowichan Bay.
The sandy-silts of Fulford Harbour probably result from
deposition from small streams, notably Fulford Creek.
Wane (1955) indicates that away from large rivers,
much of the sediment added to the inland waters is
derived from the erosion of shoreline glacial till. Most of
the eroded clay and silt fractions are carried out to sea
by the currents, leaving bottom sediments consisting of
boulders, gravel, and sand. According to WENNEKENS
(1959), erosion of shoreline glacial deposits provides a
considerable part of sediments in the San Juan Archipel-
ago. This process is undoubtedly important in the forma-
tion of bottom sediments in the Satellite Channel area
also. “Shell-gravel” from the Moresby Island grid con-
tained an assortment of small rocks (cobbles) of igneous,
sedimentary and metamorphic origin, varying in outline
from sub-rounded to sub-angular. There was also a finer
matrix containing a considerable proportion of shell frag-
ments and some intact dead bivalves enclosing a dark clay.
It seems probable that this poorly sorted sediment results
from erosion of a glacial till, possibly in situ, by selective
washing out of silt and clay, leaving behind a coarser frac-
tion, too heavy to be transported.
In the lee of the north end of Moresby Istana there
is an area relatively sheltered from the currents that sweep
through Moresby Passage and Swanson Channel. The sed-
iments here are predominantly sandy although the silt-
clay fraction of 26 samples varies from 14 to 46%. The
bathymetry of this locale suggests eddies may build up
when tidal currents reach their peak in the adjacent pas-
sages, carrying in and depositing material from suspension
Vol. 12; No. 2
in a haphazard fashion. The bottom of this sedimentary
bank slopes down to the channel trough more slowly than
the bottom in the exposed areas of Swanson Channel, to
the east. Dense bivalve populations, including large a-
dults, whose growth rings show they are 8 or more years
old, indicate the sediments of the bank are relatively
stable.
METHODS
A grid of stations was established off the north end of
Moresby Island and sampled in the summer of 1967. A
0.1m? Van Veen grab, tried initially, would not work on
“shell-gravels” and was replaced with a 52.0 1 capacity
anchor dredge, based on that in Hotme (1964). The
anchor dredge was backed by a steel plate instead of a
net, preventing “washing-out” of the sample, except at
the bucket surface. Two samples were taken at each station
and sample sizes ranged from 28.8 to 52.01 (Table 1a, 1b).
The dredge, when weighted as in Figure 5, was estimated
to penetrate to a mean depth of 7.5cm. Since sample
volumes were known, approximate sample areas could be
estimated.
In the Satellite Channel survey, 10 to 16 samples were
taken at each station using 0.1 and 0.2m” Van Veen grabs
(Table 1c).
Figure 5
The weighted anchor dredge
THE VELIGER
Page 211
Sediment samples were taken for particle analysis by a
modification of the hydrometer technique of Bouyoucos
(1951) and data are recorded in Table 1. After sample
temperatures and volumes were measured, the entire
samples were washed through a 2 mm square mesh wire
screen and all macrobenthos picked out and preserved in
10% neutral formalin. Macoma count and weight data
are recorded in DunniLx (1968) and are shown graphic-
ally in Figures 6 and 7.
The “average size” of a population of a given species
(the ratio of wet weight to number of individuals per
meter squared), assuming the full size range of the species
at that locality is sampled, is of some value as an indicator
of environmental suitability. Where “average size” is rela-
tively high for a species, the environment must be suffi-
ciently favourable to allow continued survival of individ-
uals to a (relatively) large size. Where “average sizes” at
a given locality are very small, first impressions are that
environmental conditions there are unfavourable to the
species, which is either failing to survive to a large size
(i. e., is very short lived) or has a very slow growth rate.
However, the investigator might be sampling a young pop-
ulation in a favourable environment. Counting growth
rings would show the actual age of the population. A
second sampling of a young population after the lapse of
a year or two would indicate whether or not that popula-
tion of the species was surviving and environmental con-
ditions were favourable.
Successful migration of adult specimens of Macoma
over any appreciable distance would seem unlikely, at
least in Satellite Channel, since the area supports con-
siderable numbers of bottom fish.
RESULTS
Macoma in Relation to the Physical Environment
Eight species of Macoma were found sub-littorally on the
Moresby Island: grid and two additional species, M. in-
quinata and M. nasuta, were found intertidally at the
north end of the island. Thus at least 10 of the 13 species
found in southern British Columbia (DUNNILL & ELLIs,
1969) occur within a very small area at the north end of
Moresby Island.
Macoma inquinata and M. nasuta are both shallow
water species in the study area and were not found in
grab or dredge samples either off Moresby Island or in
Satellite Channel. They will not be considered further in
this study.
Macoma yoldiformis CarPENTER, 1864 was found at a
single location, station 19 (29 m, 25.0% silt-clay) . Accord-
ding to Bernarp (1967), the species ranges from inter-
tidal to 18 m in British Columbia. It was found in Sooke
Page 212 THE VELIGER Vol. 12; No. 2
Table 1
Station List, Sample Sizes, Sample Areas
and Environmental Data
(a) Moresby Island grid - excluding stations
on “shell-gravel”
: :
6) S gD z
§ g a ag Ba 3
qe
n QA n 23) q ae SS
1 80 52.0 0.69 - 38.0
6 65 52.0 0.69 - 23.0
10 73.5 45.6 0.60 - 41.5
11 55.5 46.5 0.62 9.8 14.0
13 35.5 37.5 0.50 9.9 29.0
14 26.5 41.1 0.55 10.1 32.0
15 77.5 52.0 0.69 - 39.5
16 62.5 44.8 0.60 9.5 32.0
17 53.5 40.2 0.54 9.6 36.0
19 29.0 52.0 0.69 9.9 25.0
20 85.0 52.0 0.69 - 22.5
21 84.0 44.8 0.60 9.4 32.0
22 74.0 44.8 0.60 9.9 45.5
(b) Moresby Island grid - stations on “shell-gravel”
5
— = a iS a
g Sy 7 eS &
gS = 2 its} Si g
g c= 2. 2a. = if
Ss 3 5 ae ss S
n QA n rea} 6 ne Ss
2 55.0 49.2 0.65 10.0 42.0:
4 33.5 34.9 0.46 = 34.5
7 BLS) 52.0 0.69 9.5 43.0:
9 40.5 44.7 0.59 9.8 37.5%
" % silt-elay in sediment fraction passing a 2 mm screen
(c) Satellite Channel
€ Ss of 3 z
= oS eee ues 3
3 a. & 6 oz s
Station a =| 3 g 33 ig
A n ne FQ 2 S
FE (Fulford Harbour) 22.7 33.9! - - 69.2
CEN (Satellite Channel Center) 74.5 10.0 8.3-9.2 29.40-32.48 40.4
XS (Cross Section South) 9.0 5-33 - - 9.5
XN (Cross Section North) 9.0 1-33 - - 10.8
B (Bosun Bank) 14.9 3.1 7.4 29.90 15.3
Cc (Cowichan Bay) 61.0 12.0 8.1 29.99 95.4
W (Satellite Channel West) 68.9 10.9 8.1 29.77 82.2
E (Satellite Channel East) 50.5 5.9 7.9 30.03 25.7
‘ benthos sampled with a 0.2 m? Van Veen grab (all other stations
sampled with 0.1 m? grab)
Vol. 12; No. 2
THE VELIGER
Page 213
Basin, near Victoria (DUNNILL & Extis, 1969) at 22m
in a black silt (78.9% silt-clay), hence is tolerant of a
considerable range of sediment grain sizes. Probably most
of the stations sampled off Moresby Island and in Satellite
Channel were below the bathymetric range of M. yoldi-
formis in southern British Columbia.
Both station 19, in the lee of Moresby Island, and
Sooke Basin are in sheltered locations. The species also
occurs in a protected situation in Nanoose Bay, Vancouver
Island. Shelter from strong currents appears to be a pre-
requisite for this small, fragile species to become estab-
lished.
Despite the wide variation in sediment types found
in the survey areas (see section on Geology), few clear-cut
differences were found in the distribution of the remaining
7 species, Macoma alaskana, M. brota, M. calcarea, M.
carlottensis, M. elimata, M. incongrua, and M. lipara.
Often many of these species occurred together (Figures 6
and 7) at the same station. All are sub-littoral, with
bathymetric ranges in southern British Columbia extend-
ing below the deepest station sampled. Ranges of these
species are listed in Table 2 and bathymetric distribution
trends shown in Table 3.
Macoma alaskana Dat, 1900 was somewhat restricted
in its distribution. Where it was found, “average sizes”
of individuals tended to be greatest at the deeper stations,
below about 50m, although the greatest density was at
a shallow station (no. 19, 29m).
Like Macoma carlottensis, M. alaskana was present at
some stations and absent at others where depths and
sediments were almost identical, indicating some other
factor was limiting. It was not found at the more exposed
stations adjacent to Swanson Channel (Figure 2) where
currents might be too strong for small species such as M.
alaskana to become established, nor was it found in the
very fine-grained sediments in Cowichan Bay and Fulford
Harbour (Figure 4).
Macoma brota Dau, 1916 was widely distributed in the
survey areas, with a trend for greatest biomasses and
“average sizes” of individuals to be found at depths of
more than 40 m. Densities, biomasses and “average sizes”
were generally greater off Moresby Island and at Satellite
Channel East than elsewhere in Satellite Channel. “Aver-
age sizes” of M. brota on very fine-grained sediments were
relatively low in comparison. with those off Moresby
Island, yet would still seem too high for it to be said that
these sediments were unfavourable to the species.
Macoma calcarea (GMELIN, 1791) was present in
collections from most stations off Moresby Island, but
was not found elsewhere in Satellite Channel, although
it is known to occur a few miles north of Cowichan Bay.
Densities, biomasses, and “average sizes” all were markedly
greater in water deeper than about 40 m.
The apparent absence of Macoma calcarea in areas
where sediments were very fine-grained, i.e., Fulford
Harbour and Cowichan Bay, suggest grain size may be
an important limiting factor for the species. Possibly the
sheltered conditions in some of these localities, reflected
by fine bottom sediments, or even the annual deposition
of fine-grained material from winter run-off of the Cowi-
chan River, may also be unfavourable.
Macoma carlottensis WHITEAVES, 1880 was widely dis-
tributed in the survey area and showed no obvious pref-
erence for any particular depth zone or sediment grade.
Densities and biomasses were roughly equally great at
extremes of depth and sediment types surveyed, but “‘av-
erage sizes” tended to be low on the coarse and fine
sediment extremes.
The species was found in equally high densities in
sheltered environments in Fulford Harbour and Cowichan
Bay and in more exposed conditions at Satellite Channel
East and West.
Macoma elimata DUNNILL & Coan, 1968, the most
widely distributed Macoma in the survey area, was found
in greatest densities at shallow stations (less than 40m),
but biomasses and “average sizes” tended to be greatest
in water deeper than 40 - 50 m.
Very fine sediments, as in Cowichan Bay and at Satellite
Channel West, appear unfavourable to the species. Bio-
masses and “average sizes” there were low, even in water
deeper than 50 m. However, the species was present in
fine sediments in Fulford Harbour with relatively high
density, biomass, and “average weight.” The species ap-
parently has a tolerance for a fairly wide range of
sediment types.
Macoma incongrua (vON MarTENS, 1865) was the only
Macoma regularly found on “shell-gravels,”’ at stations
exposed to strong currents (2 - 3 knots) sweeping through
Moresby Passage. However, the greatest “‘average sizes”
of individuals of the species were found at stations away
from the “shell-gravels” where current velocities were
lower (1 - 2 knots).
At the mouth of Moresby Passage (Figure 2), a reef
with an average depth of about 36m (and exposed at
one point as Canoe Rock) affords stations below this
depth some protection from currents. Densities, biomasses,
and “average sizes” of the species on the “shell-gravels”
here increased as depth and degree of shelter increased.
At all stations surveyed, densities, biomasses, and “aver-
age sizes” of individuals tended to be greatest in water
deeper than 40-50 m.
Macoma incongrua was generally not found at stations
where sediment contained more than about 50% silt-clay
Page 214
THE
VELIGER
Vol. 12; No. 2
Table 2
Ranges in Distribution of Macoma off Moresby Island
and in Satellite Channel
Species
Macoma alaskana
Macoma brota
Macoma calcarea
Macoma carlottensis
Macoma elimata
Macoma incongrua
Macoma lipara
(m)
Depth Range
where found
14.2-65.0
14.2-85.0
29.0-85.0
9.0-80.0
9.0-85.0
9.0-85.0
29.0-85.0
[data from DuNNILL, 1968]
and where it did occur in such sediments, densities, bio-
masses, and “average sizes” were low.
Macoma lipara Daut, 1916 was collected at every sta-
tion off Moresby Island, but elsewhere was found only at
Satellite Channel East. There was a trend for densities,
E
> —
ase 2 gf
ae ¢ gS Fe
°
Ss Se) SI
14.0-36.0 18.1 11.2
15.3-95.4 52.2 449.0
14.0-45.5 225.0 873.0
9.5-95.4 344.0 13.6
9,5-95.4 157.0 117.0
9.5-82.2 89.2 182.1
14.0-45.5 17.0 392.0
Range of
“Average
Sizes” (g)
0.03- 1.2
0.03-12.0
0.7 - 7.4
0.01- 0.25
0.13- 3.0
0.02- 4.3
6.4-42.9
biomasses, and “average sizes” to be greatest at depths
of more than 40 m. “Average sizes” were generally high,
Table 3
even for this, the largest sub-littoral Macoma species in
southern British Columbia and indicate a scarcity of
young individuals. Possibly the species may have had poor
Trends in Distribution of Macoma off Moresby Island
and in Satellite Channel
Species
Macoma alaskana
Macoma brota
Macoma calcarea
Macoma carlottensis
Macoma elimata
Macoma incongrua
Macoma lipara
Maximum
Densities
Generally shallower
than 50 meters
Generally deeper
than 40 meters
Generally deeper
than 40 meters
Roughly equal at
shallow and deep
stations
Shallower than
50 meters
Roughly equal at
shallow and deep
stations
Generally deeper
than 40 meters
Maximum
Biomasses
Generally shallower
than 50 meters
Generally deeper
than 40 meters
Generally deeper
than 40 meters
Roughly equal at
shallow and deep
stations
Roughly equal at
shallow and deep
stations
Roughly equal at
shallow and deep
stations
Generally deeper
than 40 meters
Maximum
“Average Sizes”
Deeper than
40 meters
Deeper than
40 meters
Deeper than
40 meters
Roughly equal at
shallow and deep
stations
Deeper than
40 meters
At the deeper
stations
Deeper than
40 meters
Vol. 12; No. 2
THE VELIGER
Page 215
success in reproducing itself in the last few years at the
north end of Moresby Island.
The apparent absence of the species in sheltered loca-
tions and on fine-grained sediments indicates these are
unfavourable to Macoma lipara. Large adult specimens
were regularly found near the surface of the sediments,
suggesting the species is a shallow burrower. However,
PamatmMat (1961) found distribution of M. secta, another
large (intertidal) species independent of size, and this
may also be true of M. lipara.
}-—_——————___ Moresby Island
fe
20
10
S
Macoma alaskana 0 iS]
100
50
Macoma brota 0 oy = =
250
125
Macoma calcarea cm 0 Sao =
E 400
oO
a.
Z 200
5 ~
Macoma carlottensis = 0 =
= 200
A
100 Ng
NN
Macoma elimata oF S N iI
100
50
Macoma incongrua
Macoma lipara
Silty-sands
We
ZZ
DISCUSSION
In shell morphology, Macoma brota, M. calcarea, M.
elimata, and M. lipara are very similar and their similar-
ity is apparently reflected in their environmental require-
ments. They are frequently found together off Moresby
Island (Figures 6 and 7). Examination of Figure 8 shows
these species all tend to have maximum biomasses and
“average sizes” in water deeper than about 40 m and in
sediments with a silt-clay content between roughly 15 and
a
Satellite Channel
—
Shell
4
gravels
ao ttn,
~
SS
x
NS
SS
SN
~
AS
AS
AS
RS
S
SS
N
WLLLLLB
7/,]
WITT.
1/112
So 8 soa Sa
N
XN
na tae Gag =
N
S N
NS - NS
IS FS ~
SBS AS
SN 8 N SS
NOSES I RS
S NN 8 N A
S NS 8 S §
a= S&S = SS & oo = =— =
=~ SN WN
S : SS
S S § __sNaé S S
U1.
W772
—_
W FCENE B XN XCXS
2 a a QO C
Station
Figure 6
Densities of populations of sub-littoral species of Macoma
off Moresby Island and in Satellite Channel
Page 216 THE VELIGER i Vol. 12; No. 2
50%. Macoma calcarea and M. lipara are apparently able situations, settle there but are then short-lived and
more restricted in their distribution than M. brota and M. probably fail to survive to reproductive maturity, in which
elimata and perhaps are less tolerant of the finer grained case they are dependent on replacement from outside
sediments. populations in more favourable locations.
The presence of Macoma brota and M. elimata on these The smaller species, Macoma alaskana and M. incon-
very fine-grained sediments (containing more than 80% grua (Figure 9) also appear to favour water deeper than
silt-clay) and the low “average size” of individuals, sug- 40-50m and sandy sediments. Macoma carlottensis is
gest the young of these animals are carried into unfavour- exceptional and appears tolerant of shallower water and
Moresby Island ————————+}— Satellite Channel —of
|>—_——_———_ Silty- —— :
ilty-sands |— er
10 <
SN
Macoma alaskana S
N
S
500 0 B = ay a —)
250
Macoma brota
~
0 1000 P S --8s - ee
oO
i=
©
A 500
Macoma calcarea =
= ~ ©
2 20 0 = = = oo_ 2 SS S =
~_
o
a ~“
= 10 3 K
wo S S Ss
. ~ N ~
Macoma carlottensis S S S N :
g S § 2 NSS SN
fa) 0 900 = = — SS A oso IN IS
S
Macoma elimata N S SNANaWAN A N S
N NSNNNSASAAB N ~ N
200 0 SS SSSSSSNSSHSS8S -~-anNnrwtWNS SS
100
Macoma incongrua
S
250 N
N
Macoma lipara S
S
0
ne
122 2 47 9 G W FCENE B XNXCXS
Station
Figure 7
Biomasses of populations of sub-littoral species of Macoma
off Moresby Island and in Satellite Channel
Wola l2-.INo. 2
Macoma brota
THE VELIGER
Page 217
Macoma elimata
2
= 1
a
3
a=)
EB
a=)
: LD AT SO ae
5 ®
= 10
a)
: Macoma lipara
ty
a 30
d Bal wet weight/m* (g)
: © ei @ depth (m) 0-10 10-100 100-1000
~ ® jo O° ©
& 5-50 9% 8
10 50- 75 ®@ ® e
CO es 75-100 & a
e 20 40. 60 80. 100
% Silt-Clay
Figure 8
Plots of “average size” of Macoma brota, Macoma calcarea,
Macoma elimata and Macoma lipara versus depth and % silt-clay
in sediments
a greater range of sediment types than the other species
examined.
The mean number of different species of Macoma oc-
curring in hauls off the north end of Moresby Island on
silty-sands was 5.3 compared to only 1.7 on “shell-gravel”
(Figures 6 and 7). In the rest of Satellite Channel, the
mean was 3.3. The paucity of Macoma species found on
“shell-gravels” seems to result from the prevailing strong
currents there which may prevent them from becoming
established or may hinder their feeding. The siphons of
species such as Modiolus modiolus (Linnagus, 1758) and
Humilaria kennerleyi (Reeve, 1863), which flourish on
the “shell-gravels,” are very short and have wide bores,
in diametric contrast to those of Macoma, which are very
long and of small bore. However, observations made on
sub-littoral Macoma species in aquaria suggest some of
these are not obligatory detrital feeders but can shorten
their inhalant siphons and feed from the water above the
bottom. This behaviour has also been reported for the
Atlantic species, Macoma balthica (Linnazus, 1758)
(BRAEFIELD & NEWELL, 1961).
The mean wet weight/m’ of all macomas taken with the
0.1 m? Van Veen grab from 8 stations below 40 m off the
north end of Moresby Island, was 1737 g (= 114g dry
organic matter/m’, using the mean of the conversion fac-
tors for Macoma listed in THorson (1957) ). It is evident
that the area has an exceptionally rich infaunal standing
crop. In comparison, Hote (1963) and Sanpers (1956)
obtained total faunal mean dry organic matter weights of
11.2 and 54.627 g/m? for the English Channel and Long
Island Sound, respectively. Food must be plentiful off
Moresby Island to support this large standing crop and
may not at present be a limiting factor governing the den-
sities and biomasses of Macoma in this location.
Page 218
THE VELIGER
Vol. 12; No. 2
1.5 Macoma alaskana
0 20 40 60 80 100
0.3 Macoma carlottensis
Mean Wet Weight per Individual (g)
Macoma incongrua
wet weight/m? (g)
0-10 10-100 100 - 1000
depth (m)0-5 5-10 10-15
0. & 2 ©
Bo 8 fF
50-75 ¢ ©@ 8
75-100 8 a ie
% Silt-Clay
Figure 9
Plots of “average size” of Macoma alaskana, Macoma carlottensis
and Macoma incongrua versus depth and % silt-clay in sediments
Frequently, 6 different species of Macoma were taken
in a single anchor dredge haul at the north end of Moresby
Island, from an estimated area of less than 0.7m’, and 5
different species were taken from areas of 0.1m? with the
Van Veen grab (see DuNNILL, 1968). At station 19, the
two anchor dredge hauls captured between them all 8
species of Macoma found sub-littorally in the survey
(Figures 6 and 7). Macoma yoldiformis, collected at sta-
tion 19, is not figured since the authors consider it a
littoral species.
According to Gause (1934), as a result of competition,
two similar species scarcely ever occupy similar niches (by
niche he means “. . . what place the given species oc-
cupies in a community, i. e. what are its habits, food and
mode of life.”), but displace each other so that each
utilizes “certain peculiar kinds of food and modes of life
in which it has an advantage over its competitor.”
It seems probable that the merging and diverging of
currents flowing through the channels at the north end of
Moresby Island would generate extensive eddies. These
might aid in dispersal of larvae of all local species over
the submerged bank and could thus account for the dif-
ferent species being brought together initially. There is
some evidence (see above) that food may be more than
adequate to support the existing Macoma populations off
Moresby Island so that interspecific competition is lessened,
thus permitting the coexistence of these closely related
species. However, determination of the exact “niches”
occupied by each of these coexisting species of Macoma
off Moresby Island is beyond the scope of the present
study.
LITERATURE CITED
AppicotT, WARREN OLIVER
1952. Ecological and natural history studies of the pelecypod
genus Macoma in Elkhorn Slough, California. Master of
Arts thesis, Stanford Univ., 89 pp.
BERNARD, FRANK a
1967. | Prodrome for a distributional check-list and bibliogra-
phy of the marine Mollusca of the west coast of Canada.
Fish. Res. Brd. Canada, Tech. Rept. No. 2: 261 pp.
Bouyoucos, GrorcE JOHN
1951. A recalibration of the hydrometer method for making
mechanical analyses of soils. Agron. Journ. 43: 434 - 438
BRAEFIELD, A. E. « G. E. NEWELL
1961. The behavior of Macoma balthica (L.)
Biol. Assoc. U. K. 41: 81 - 87
Journ. Mar.
Vol. 12; No. 2
THE VELIGER
Page 219
DuNNILL, Ropert M.
1968. A taxonomie and ecological investigation of the genus
Macoma (Pelecypoda) in southern British Columbia.
Master of Sci. thesis, Univ. Victoria, 155 pp.
DuNNILL, Rospert M. & EUGENE Victor CoAN
1968. A new species of the genus Macoma (Pelecypoda) from
west American coastal waters, with comments on Macoma cal-
carea (GMELIN) 1791. National Mus. Canada, Nat. Hist.
Paper 43: 19 pp.
DunnILL, Rosert M. & Derek V. ELLIS
1969. Recent species of the genus Macoma (Pelecypoda) in
British Columbia. National Mus. Canada, Nat. Hist. Pap.
45: 34 pp.
Exuis, DEREK V.
1967. Quantitative benthic investigations. IT. Satellite Channel
species data, February 1965 - May 1967. Fish. Res. Brd.
Canada, Tech. Reprt. No. 35: 8 pp.; 169 tables; 2 figs.
in press Quantitative benthic investigations. III. Locality and
environmental data for selected stations (mainly from Satellite
Channel, Straits of Georgia and adjacent inlets). Fish. Res.
Brd. Canada, Tech. Rprt.
Gause. G. F
1934. The struggle for existence. (Repr. 1964) Hafner, New
York, 163 pp.
HER.LINVEAUxX, Ricuarp H.
1962. | Oceanography of Saanich Inlet in Vancouver Island,
British Columbia. Journ. Fish. Res. Brd. Canada 19 (1) :
1-37
HER.LINVEAUx, RicHarp H., O. D. Kennepy «& H. J. HoLiisTEr
1960. (MS) Oceanographic data record. Coastal Seaways project,
November 16-December 11, 1959. Fish. Res. Brd. Canada,
MS Reprt. Oceanogr. & Limnol. Ser. No. 58: 134 pp.
Hotme, Norman A.
1953. The biomass of the bottom fauna in the English Channel
off Plymouth. Journ. Mar. Biol. Assoc. U. K. 32: 1 - 49
1964. | Methods of sampling the benthos. Adv. Mar. Biol.
2: 171 - 260
Jones, Norman S.
1950. Marine bottom communities. Biol. Rev. 25: 283-313
MacGinitTi£, GEorGE EBER
1935. Ecological aspects of a California marine estuary. Amer.
Midl. Natur., 16 (5) : 629 - 765
PaciFic OcEANOGRAPHIC Group
1959a. (MS) Physical and chemical data record. Coastal Sea-
ways project, March 3lst-April 22nd, 1959. Fish. Res.
Brd. Canada. MS Rprt. Oceanogr. Limnol. Ser. No. 47: [only
p. 11 seen]
1959b. (MS) Physical and chemical data record. Coastal Sea-
ways project, June 8-July 1, 1959. Fish. Res. Brd. Canada.
MS Rprt. Oceanogr. Limnol. Ser. No. 52: 210 pp.
PAMATMAT, Mario M.
1961. A study of various aspects of feeding and related prob-
lems in Macoma secta and Neoamphitrite robusta. | Unpubl.
MS Univ. Washington, 31 pp.
SANDERS, Howarp L.
1956. | Oceanography of Long Island Sound, 1952-1954. X.
Biology of marine bottom communities. Bull. Bingham
Oceanogr. Coll. 15: 345 - 414
SHELForD, Victor E. « E. T. TowLer
1925. Animal communities of the San Juan Channel and
adjacent areas. Univ. Wash. Publ., Puget Sound Biol. Sta.
5: 31-73
SHELForD, Victor E., A. O. WeesE, L. A. Rice, D. I. RasMUSSEN &
A. MacLean
1935. Some marine biotic communities of the Pacific coast of
North America. Part I. General survey of the communities --
their extent and dynamics. Ecol. Monogr. 5 (3) : 249 - 332
Tuorson, GUNNAR
1957. Bottom communities. Jn Treatise on marine ecology
and paleoecology. J. W. Hedgpeth (ed.) Geol. Soc. Amer.
Mem. 67, 1: 461 - 534
Tutty, J. P « A. J. Dopimeap
1957. Properties of the water in the Strait of Georgia, British
Columbia, and influencing factors. Journ. Fish. Res. Brd.
Canada 14 (3): 241-319
Wa.picHuck, M.
1957. Physical oceanography of the Strait of Georgia, British
Columbia. Journ. Fish. Res. Brd. Canada 14 (3) :321 - 486
Wans, Frank Fenc-Hut
1955. Recent sediments in Puget Sound and portions of Wash-
ington Sound and Lake Washington. Ph. D. thesis, Univ.
Wash., 160 pp.
WENNEKENS, Marce P.
1959. Marine-environment and macro-benthos of the waters
of Puget Sound. San Juan Archipelago, southern Georgia
Strait, and Strait of Juan de Fuca. Ph. D. thesis, Univ.
Wash., 298 pp.
Page 220
THE VELIGER
Vol. 12; No. 2
Additional Bathymetric and Locality Data
for some Opisthobranchs and an Octopus
from Santa Barbara County, California
RICHARD S. LEE
Santa Barbara Museum of Natural History, Santa Barbara, California 93105
AND
PATRICK BROPHY
Pacific Bio-Marine, Venice, California go291
RECENTLY AN EXCELLENT annotated list of the opistho-
branch mollusks from Santa Barbara County (SPHON &
Lance, 1968) summarized all the geographic and bathy-
metric information available on the local members of this
group up to the date of publication. Since that time the
junior author of this report has uncovered additional
information, including new bathymetric records and the
presence of 8 unlisted species in the waters off Santa
Barbara County.
LIST or SPECIES
Acanthodoris atrogriseata O’DonocHUE, 1927
Intertidal at Serena Cove, Carpinteria.
First record for Santa Barbara County.
Acanthodoris hudsont MacFartanp, 1905
Trawled at 700 (?) feet at Gaviota.
Found in a sunken holdfast.
First record for Santa Barbara County.
Acanthodoris nanaimoensis O’DoNoGHUE, 1921
Intertidal, Purisima Point.
Not previously recorded from Santa Barbara County,
although collected at Shell Beach, San Luis Obispo
County (RoLLer « Lone, 1969).
Anisodoris nobilis (MacFarLanp, 1905)
Intertidal to 840 feet off Santa Cruz Island.
Archidoris montereyensis Coorer, 1863
Intertidal to 840 feet off Santa Cruz Island.
Berthellina engeli GARDNER, 1936
25 feet, Santa Cruz Island. Previously recorded only
as far north as San Diego.
Cadlina flavomaculata MacFarianp, 1905
Intertidal to 723 feet off Port Hueneme (Ventura
County).
Dendrodoris sp.
Undescribed white and brown species.
30 feet, off Naples, Santa Barbara County.
Dendronotus tris Cooper, 1862
15 to 702 feet, off Gaviota.
Doto wara Marcus, 1961
Santa Barbara Yacht Harbor.
First record for Santa Barbara County.
Gastropteron pacificum Brercu, 1894
Intertidal to 804 feet. The first recorded specimen
from the County was taken from the backwash of the
filtration unit of the Santa Barbara Undersea Gar-
dens, while the remaining records are based on speci-
mens trawled at depths of 510 to 840 feet from
Rincon Point and Gaviota (BerTscH, 1969).
Hermaeina oliviae MAcFaRLAND, 1966
Holdfast, Santa Barbara Channel. Depth unknown.
First record for Santa Barbara County.
Petelodoris spongicola MAcFaRLAND, 1966
One specimen taken in 690 feet in a sunken kelp
holdfast off Gaviota. Not previously recorded from
Santa Barbara County.
Phidiana pugnax Lance, 1962
Intertidal to 702 feet, off Gaviota, found in a sunken
kelp holdfast.
Philine bakeri Dau, 1919
510 to 572 feet, off Gaviota.
Phyllaplysia taylori Dai, 1900
15 feet, Santa Cruz Island.
Not previously recorded from Santa Barbara County.
Pleurobranchaea californica MacFaRLAND, 1966
Subtidal to 714 feet, off Gaviota.
Vol. 12; No. 2
In October and November of 1968, 3 female Opistho-
teuthis californiana Berry, 1949 were trawled from off
the Santa Barbara coast. The first was taken on October
24, 1968, at approximately 840 feet, off Gaviota; the
second, gravid, was taken at the same depth off Santa
Barbara Harbor. Both of these specimens were collected
by the junior author. The third specimen was taken at a
similar depth by Mr. Harold Durrah, a commercial
fisherman, who kindly brought it to our attention.
Berry (1949) originally described the species on the
basis of 2 females taken off Eureka, Humboldt County,
California, and then later (1955) described a male from
the same area. A substantial range extension southwards
to Monterey was noted by Puitups (1961). On the
basis of these notes, we feel that the Santa Barbara speci-
mens represent the southernmost records for this unusual
webbed octopus. However, it is suspected that as commer-
cial fishermen continue their deep water exploration of
this coast, farther southern records will be established.
ACKNOWLEDGMENTS
We wish to thank Mrs. Thomas Rogers of the Santa
Barbara Museum of Natural History for her aid in the
identification of the opisthobranchs; Mr. Allyn G. Smith
THE VELIGER
Page 221
of the California Academy of Sciences for his encourage-
ment and suggestions on this note; Mr. Harold Durrah,
Mr. Shane Anderson, Mr. Samuel Spaulding of Santa
Barbara for supplying specimens and data.
LITERATURE CITED
Berry, SAMUEL STILLMAN
1949. <A new Opisthoteuthis from the eastern Pacific.
Leafl. Malacol. 1 (6): 23 - 26
1955. The male flapjack devilfish.
41 (3): 219-224
BertscH, Hans
1969. A note on the range of Gastropteron pacificum (Opistho-
branchia: Cephalaspidea) . The Veliger 11 (4): 431 - 433
(1 April 1969)
Calif. Fish & Game
Puituips, Juuius B.
1961. Two unusual cephalopods taken near Monterey.
Calif. Fish & Game 47 (4) :416 - 417
Router, RicHarp A. & STEVEN J. Lone
1969. An annotated list of opisthobranchs from San Luis Obis-
po County, California. The Veliger 11 (4): 424-430; 1
lap (1 April 1969)
SpHON, GALE G., Jr. & JAMES Ropert LANCE
1968. An annotated list of nudibranchs and their allies from
Santa Barbara County, California. Proc. Calif. Acad.
Sci. 36 (3): 73 - 84; 1 fig. (25 September 1968)
Page 222
THE VELIGER
Vol. 12; No. 2
A Bibliography of the Biological Writings
of Puitie PEARSALL CARPENTER
EUGENE V. COAN
Department of Geology, California Academy of Sciences,
THE FOLLOWING BIBLIOGRAPHY has been prepared and is
presented here for several reasons. First, CARPENTER was
the most important early worker on the molluscan fauna
of the eastern Pacific, and a complete bibliography of his
papers is nowhere available. Secondly, it has been possible
to date certain papers more accurately than in the past.
I have relied on RHEES (1882) for the dates of the publi-
cations of the Smithsonian Institution, on NoLan (1913)
for the Proceedings of the Academy of Natural Sciences of
Philadelphia, on FiscHEr-Pretre (1937) for the Journal
de Conchyliologie, and on Duncan (1937) for the Pro-
ceedings of the Zoological Society of London. Other
sources of dates are cited in the bibliography. Finally, I
have listed a number of papers omitted from previous
bibliographies. While none of these papers contains de-
scriptions of new taxa, they do have important discussions.
CARPENTER made contributions to the concepts of species
and variation within species, to nomenclatural, systematic,
and curatorial techniques, and to biogeography. At a time
when the nomenclatural stability of the West American
fauna seems assured, workers are becoming more interested
in such corollary studies. These smaller papers of his are
of significance to the development of ideas on these topics.
Bibliography of CaRPENTER’s Papers
1855. List of four hundred and forty species of shells from
Mazatlan. Brit. Assoc. Adv. Sci., Rept. 24 (for 1854)
[Trans.]: 107 - 108
1856. Descriptions of (supposed) new species and varieties of
shells, from the Californian and west Mexican coasts, principal-
ly in the collection of Hugh Cuming, Esq. Proc. Zool. Soc.
London (for 1855) [pt. 23] (298): 228-232 (5 February) ;
(299) : 233 - 235 (23 February)
1856. Notes on the species of Hipponyx inhabiting the Ameri-
can coasts, with descriptions of new species. Proc. Zool.
Soc. London (for 1856) [pt. 24] (301): 3-5 (16 June)
Golden Gate Park, San Francisco, California 94118
1856. Description of new species of shells collected by Mr. T.
Bridges in the Bay of Panama and its vicinity, in the collection
of Hugh Cuming, Esq. Proc. Zool. Soc. London (for
1856) [pt. 24] (310): 159-160; (311): 161-166 (11 Nov.)
1856. Description of new species and varieties of Calyptrae-
idae, Trochidae, and Pyramidellidae, principally in the col-
lection of Hugh Cuming, Esq. Proc. Zool. Soc. London
(for 1856) [pt. 24] (311): 166-171 (11 November)
1857. see GouLp & CarPENTER (1857) (7 Jan.)
1857. Monograph of the shells collected by T. Nuttall, Esq.,
on the Californian coast, in the years 1834-5. Proc. Zool.
Soc. London (for 1856) [pt. 24] (314): 209-224; (315):
225 - 229 (26 January 1857)
1857. First steps towards a monograph of the Recent species of
Petaloconchus, a genus of Vermetidae. Proc. Zool. Soc.
London (for 1856) [pt. 24] (320): 313-317; 8 text figs.
(10 March 1857)
1857. | Report on the present state of our knowledge with re-
gard to the Mollusca of the west coast of North America.
Brit. Assoc. Adv. Sci., Rept. 26 (for 1856): 159-368+4 pp.;
plts. 6-9 (pre 22 April) [dating: the following, p. iv]
1857. | Catalogue of the collection of Mazatlan shells, in the
British Museum: collected by Frederick Reigen, ... London
(British Museum) i - iv+ix - xvi+552 pp. (1 August) [dating:
SHERBORN, 1934] [reprinted: Paleo. Res. Inst., Ithaca, New
York, 1967]
1857. Catalogue of the Reigen collection of Mazatlan Mollus-
Warrington (Oberlin) i - viii+
i-xii+552 pp. (1 August) [dating: CarpENTER, 1872, p. xi;
TrEDALE, 1916, p. 36]
ca, in the British Museum.
1858. Note on peculiarities of growth in Caecidae. Brit.
Assoc. Adv. Sci., Rept. 27 (for 1857) [Trans.]: 102
1858-1859. First steps towards a monograph of the Caecidae, a
a family of rostiferous Gastropoda. Proc. Zool. Soc. Lon-
don (for 1858) [pt. 26] (372): 413-416; (373): 417-432
(14 December 1858) ; (374) : 433 - 444 (“Jan. to May” 1859)
1860. Notice of the shells collected by Mr. J. Xantus, at Cape
St. Lucas. Acad. Nat. Sci. Philadelphia, Proc. (for 1859)
11 (21 - 23): 331 - 332 (12 January)
Vol. 12; No. 2
1860. Catalogue of the Reigen collection of Mazatlan Mol-
lusca, presented to the State Cabinet ..., being the first dupli-
cate of the collection presented to the British Museum. — Ann.
Rept. (Regents Univ. State of New York) on the condition of
the State Cabinet Nat. Hist. 13 (for 1859) [Appendix
D]: 21-36 (post 10 April) [State Senate Doc. 89]
1860. Check lists of the shells of North America: No. 1 (a).
West Coast: Oregonian and Californian province. 4 pp. No. 2
(b) : West Coast: Mexican and Panamic province. 13 pp.
Smithson. Inst. Misc. Coll. 2 (128) [art. 6]: 44 pp. by several
authors (June)
1860. Lectures on the shells of the Gulf of California.
Smithson. Inst. Ann. Rept. (for 1859): 195-219; 6 text figs.
(post 14 June)
1861. On the progress of natural science in the United States
and Canada [chiefly a discussion of the Smithsonian Institu-
tion]. Brit. Assoc. Adv. Sci., Rept. 30 (for 1860) [Trans.]: 109
1861. Lectures on Mollusca; or “‘shell-fish” and their allies.
Smithson. Inst. Ann. Rept. (for 1860) (152): 151-283 [also
issued separately, 140 pp.]
1862. On the cosmopolitan operations of the Smithsonian In-
stitution. British Assoc. Adv. Sci., Rept. 31 (for 1861)
[Trans.]: 137
1862. On the variations of Tecturella grandis. Brit. Assoc.
Adv. Sci., Rept. 31 (for 1861) [Trans.]: 137
1864. Review of Prof. C. B. Adams’s ‘Catalogue of the Shells
of Panama,’ from the type specimens. Proc. Zool. Soc. Lon-
don (for 1863) (3): 339-369 (April) [reprinted in Car-
PENTER, 1872 (B): 173-205 (also pp. 1 - 31)]
1864. | Diagnoses of new forms of mollusks collected at Cape
St. Lucas by Mr. Xantus. Ann. Mag. Nat. Hist. (3) 13
(76): 311-315 (April); (78): 474-479 (June); 14 (79):
45-49 (July) [reprinted in Carpenter, 1872 (C): 207 - 221
(also pp. 1 - 13) ; first installment discussed by REEvE, 1864]
1864-1866. Descriptions of new marine shells from the coast of
California. Parts I-III. Calif. Acad. Sci., Proc. 3: 155 - 159
(July 1864) ; 175-176 (Dec. 1864) ; 177 (Jan. 1865) ; 207 to
208 (post 4 September 1865) ; 209 - 224 (February 1866)
1864. | Supplementary report on the present state of our know-
ledge with regard to the Mollusca of the west coast of North
America. Brit. Assoc. Adv. Sci., Rept. 33 (for 1863) :
517 - 686 (post 1 August) [reprinted in CarPEeNnTER, 1872 (A):
1 - 172]
1864. On the present state of malacological nomenclature.
Ann. Mag. Nat. Hist. (3) 14 (80): 155 - 158 (August)
1864-1865. Diagnoses of new forms of Mollusca from the Van-
couver District. Ann. Mag. Nat. Hist. (3) 14 (84): 423
to 429 (Dec. 1864); 15 (85): 28-32 (Jan. 1865) [reprinted
in CaRPENTER, 1872 (E): 233 - 246 (also pp. 1 - 12)]
1865. | Diagnoses of new forms of Mollusca from the west coast
of North America, first collected by Col. E. Jewett. Ann.
Mag. Nat. Hist. (3) 15 (87): 177-182 (March); (89):
394 - 399 (May) [reprinted in CarpENnTER, 1872 (K): 277 - 289]
THE VELIGER
Page 223
1865. Diagnoses de mollusques nouveaux provenant de Cali-
fornie et faisant partie du Musée de I’ Institution Smithsonienne.
Journ. Conchyl. 13 [(3) 5] (2): 129-149 (4 April) [reprinted
in Carpenter, 1872 (M): 295 - 317]
1865. | On the connection between the Crag formations and the
Recent North Pacific faunas. Geol. Mag. 2 (10): 152 - 154
(April 1865)
1865. On the Pleistocene fossils collected by Col. E. Jewett at
Sta. Barbara (California) ; with descriptions of new species.
Ann. Mag. Nat. Hist. (3) 17 (100): 274-278 (April) [re-
printed in Carpenter, 1872 (N): 319-325, and in Da tt,
1909 (XII): 189-191]
1865. | Contributions towards a monograph of the Pandoridae.
Proc. Zool. Soc. London (for 1864) (3): 596-603 (May) [re-
printed in Carpenter, 1872 (D): 223 - 232]
1865. Diagnoses of new forms of Mollusca collected by Col. E.
Jewett on the west tropical shores of North America. Ann.
Mag. Nat. Hist. (3) 15 (89): 399-400 (May) [reprinted in
CarrenterR, 1872 (L): 291 - 294]
1865. Diagnoses of new forms of Mollusca from the Vancouver
District. Proc. Zool. Soc. London (for 1865) (1): 201 - 204;
4 text figs. (June) [reprinted in Carpenter, 1872 (F): 247 - 252]
1865. | Diagnoses of new species and a new genus of mollusks
from the Reigen Mazatlan collection: with an account of
additional specimens presented to the British Museum. Proc.
Zool. Soc. London (for 1865) (1): 268 - 274 (June) [reprinted
in CarPENTER, 1872 (G): 253 - 261]
1865. Descriptions of new species and varieties of Chitonidae
and Acmaeidae, from the Panama collection of the late Prof.
C. B. Adams. Proc. Zool. Soc. London (for 1865) (1):
274 - 277 (June) [reprinted in CarPEeNnTER, 1872 (H): 263 - 268]
1865. | Diagnoses of new species of mollusks, from the west
tropical region of North America, principally collected by the
Rev. J. Rowell, of San Francisco. Proc. Zool. Soc. London
(for 1865) (1): 278-282 (June) [reprinted in CarPENTER,
1872 (1): 269 - 275]
1865. Diagnoses specierum et varietatum novarum molusco-
rum [sic], prope Sinum Pugetianum a Kennerlio Doctore, nuper
decesso, collectorum. Acad. Nat. Sci. Philadelphia, Proc.
17 (2): 54-64 (7 August 1865)
1865. On the connection between the Crag formations and the
Recent North Pacific faunas. Brit. Assoc. Adv. Sci., Rept.
34 (for 1864) [Trans.]: 83
1865. Description of two species of Chitonidae, from the col-
lection of W. Harper Pease, Esq. Proc. Zool. Soc. London
(for 1865) (2): SI1-512 (October)
1865. [List of synonyms]. pp. 516-517 %n Witt1AM Harper
Pease, “Descriptions of new genera and species of marine shells
from the islands of the central Pacific.” Proc. Zool. Soc.
London (for 1865) (2): 512-517 (October) [reprinted in
Amer. Journ. Conch. 2 (4): 379-380, 6 October 1866; dis-
cussed by PEAsE, 1868]
1866. On the regard due to usage and utility, as well as mere
priority in fixing zoological nomenclature. Brit. Assoc. Adv.
Sci., Rept. 35 (for 1865) [Trans.]: 83
Page 224
1866. On the Acmaeidae of the Vancouver and Californian
province. Amer. Journ. Conch. 2 (4): 332 - 348 (6 Oct.)
1867. [Description of Zonites (Conulus) priscus}. pp. 331 - 333;
1 text fig. in JoHN Witu1aAM Dawson, “On the dis-
covery of a new pulmonate mollusk [Zonites (Conulus) priscus
Cpr.] in the Coal-formation of Nova Scotia.” Geol. Soc.
London, Quart. Journ. 23 [pt. 1] (3) : 330 - 333; 1 text fig.
1869. Catalogue of the family Pandoridae. Jn “Catalogue and
synonymy of the genera, species and varieties of Recent Mol-
lusca described prior to January 1‘, 1867, with dates of pub-
lication, references to plates, and localities. Compiled and pub-
lished under the authority of the Conchological Section of the
Academy of Natural Sciences of Philadelphia,” part 2: 69 - 71.
Amer. Journ. Conch. 4 (5) [Appendix]: 69 - 71 (6 May)
1872. [Description of Choristes]. pp. 292 - 294; plt. 7, fig. 13
in JoHN Witi1aAM Dawson, “Notes on the post-Pliocene geo-
logy of Canada: III. — Revision of post-Pliocene fossils of
Canada: Sub-kingdom Mollusca. IV [“III’’]. — General con-
clusions.” Canadian Naturalist & Quart. Journ. Sci. (new ser.)
6 (4): 369-416; plts. 4-7 (August) [plates possibly part of
6 (3) (April)]
1872. | The mollusks of western North America. Embracing the
second report made to the British Association on this subject,
with other papers; reprinted by permission, with a general
index. Smithson. Inst. Misc. Coll. 10 (252): xii+325+
13-121 pp. (December 1872)
1873. | On the generic affinities of the New England chitons.
Bull. Essex Inst. 5 (9): 152-155 (September 1873)
1874. On the generic affinities of the New-England chitons.
Ann. Mag. Nat. Hist. (4) 13 (74): 119-123 (Feb. 1874)
1876. On the primary divisions of the Chitonidae. Amer.
Assoc. Adv. Sci., Proc. 24 (for 1875) [pt. 2 — B] (3) : 236 - 237
Goutp, Aucustus Appison & PHILIP PEARSALL CARPENTER
1857. Descriptions of shells from the Gulf of California, and
the Pacific coasts of Mexico and California. Part II. Proc.
Zool. Soc. London (for 1856) [pt. 24] (313): 198 - 208
[“Part I” by Goutp alone, published elsewhere] [concerning
the species described in this paper, see JoHNSOoN, 1964]
Selected Papers about CARPENTER
and his Specimens; and Collations of Books and
Journals containing CARPENTER’s Papers
ANONYMOUS
1877. | The late Dr. Philip P Carpenter. The Montreal
Gazette (25 May 1877)
1877. Dr. Philip P. Carpenter. Nature 16 (May 31): 84
1895. [Notes concerning the purchase of part of Carpenter’s
collection by the Field Museum, Chicago.] The Nautilus
9 (3): 35 (8 July 1895); 9(5): 55 (3 September 1895)
Brann, Doris C.
1966. _ Illustrations to “Catalogue of the collection of Mazat-
lan shells” by Philip P. Carpenter. Ithaca, New York (Pa-
leo. Res. Inst.) 111 pp.; 60 plts. (1 April 1966)
THE VELIGER
Vol. 12; No. 2
CaRPENTER, RUSSELL LANT
1880. Memoirs of the life of Philip Pearsall Carpenter ...
chiefly derived from his letters. London (Paul) xiii+ 360
pp-; portrait and 5 illust.
Dai, WILLIAM HEALEY
1877. [Note concerning P. P. Carpenter.]
11 (8): 504-505
Amer. Natural.
(August 1877)
1909. Contributions to the Tertiary paleontology of the Pacific
Coast. I. The Miocene of Astoria and Coos Bay, Oregon.
U.S. Geol. Surv. Prof. Paper 59: 279 pp.; 23 plts.; 14 text figs.
[with 13 Appendices reprinting scarce papers on West Ameri-
can paleontology] (2 April 1909)
Dawson, JoHN WILLIAM
1878. [Note about Carpenter].
Journ. Sci. (2) 8 (8): 445-446
The Canad. Natur. & Quart.
(July 1878)
Duncan, EF Martin
1937. On the dates of publication of the Society’s ‘Proceedings,’
1859 - 1926. With an appendix containing the dates of publi-
cation of ‘Proceedings,’ 1830 - 1858, compiled by the late F H.
Waterhouse, and of the “Transactions, 1830 - 1869, by the late
Henry Peavot, originally published in P. Z.S. 1893, 1913.
Proc. Zool. Soc. London (for 1937) [A] (1): 71-84 (April)
FISCHER-PIETTE, EDOUARD
1937. Dates de publication du “Journal de Conchyliologie” de
1861 a 1900. Journ. de Conchyl. 81 [(4) 35] (1): 88-92
(15 April 1937)
Gatpraitn, I. C. J. & STANLEY PETER DANCE
1961. British Museum (Natural History) [a discussion of the
Mazatlan collection and Catalogue]. Am. Malacol. Union,
Ann. Rept. & Bull. (for 1960) 27: 10-12 (1 Jan. 1961)
Haas, Fritz
1944. [Letter mentioning Carpenter material in the Field Mu-
seum, Chicago.] Am. Malacol. Union, Ann. Rept. & Bull.
(for 1943): 6-7
IREDALE, Tom
1916. | On some new and old molluscan generic names. _ Proc.
Malacol. Soc. London 12 (1): 27 - 37 (20 March 1916)
Jounson, RicHarp I.
1964. | The Recent Mollusca of Augustus Addison Gould.
U.S. Nat. Mus. Bull. 239: 182 pp.; 43 plts.
KEEN, A. Myra
1968. | West American mollusk types at the British Museum
(Natural History) IV. Carpenter’s Mazatlan collection.
The Veliger 10 (4) : 389 - 439; plts. 55-59; 171 text figs.
(1 April 1968)
Noxan, Epwarp J. (Editor)
1913. An index to the scientific contents of the Journal and
Proceedings of the Academy of Natural Sciences of Phila-
delphia. xiv+ 1419 pp.
PaLMER, KATHERINE EVANGELINE Hitton (VAN WINKLE)
1921. Illustrations and descriptions of fossil Mollusca con-
tained in the paleontological collections at Cornell University.
Bull. Amer. Paleo. 8 (36): 347 - 358 (also pp. 1-12); plt. 15
(1 March 1921)
Vol. 12; No. 2
1945. Molluscan types in the Carpenter collection in the Red-
path Museum. The Nautilus 58 (3): 97-102 (19 Feb.)
1951. Catalog of the first duplicate series of the Reigen collec-
tion of Mazatlan shells in the State Museum at Albany, New
York. New York State Mus. Bull. 342: 79 pp.; 1 plt. (Jan.)
1958. ‘Type specimens of marine Mollusca described by P. P.
Carpenter from the West Coast (San Diego to British Colum-
bia) . Geol. Soc. Amer. Mem. 76: viii+376 pp.; 35 plts.
(8 December 1958)
1963. Type specimens of marine Mollusca described by P. P.
Carpenter from the west coast of Mexico and Panama. _ Bull.
Amer. Paleo. 46 (211) : 285 - 408; plts. 58 - 70 (22 Oct.)
SHERBORN, CHARLES DAVIES
THE VELIGER
Page 225
PEASE, WILLIAM HARPER
1868. Notes on the remarks of Dr. P. P Carpenter (published
in Proc. Zool. Soc. Lond., 1865) on certain species of marine
Gastropoda, named by W. Harper Pease. Amer. Journ.
Conch. 3 (3): 231 - 234 (2 January 1868)
REEVE, LoveLL AuGusTus
Ann. Mag. Nat. Hist.
(May 1864)
1864. “New forms of mollusks?”
(3) 13 (77): 440
RueEEs, WILLIAM J.
1882. Catalogue of publications of the Smithsonian Institu-
tion (1846 - 1882), with an alphabetical index of articles ...
Smithson. Inst. Misc. Coll. 27 (478) : xiv+328 pp.
1934. Dates of publication of catalogues of natural history
(post 1850) issued by the British Museum.
Hist. (10) 13 (74): 308 - 312
STEARNS, Rospert Epwarps CARTER
on the late Doctor Philip P Carpenter,
1877. Remarks ...
Ann. Mag. Nat.
(February 1934)
before the California Academy of Sciences, July 2d, 1877.
Calif. Acad. Sci. 5 pp.
Page 226
NOTES & NEWS
Russian Contributions to Malacology
BY
KENNETH J. BOSS
Museum of Comparative Zoology, Harvard University
Cambridge, Massachusetts 02138
RUSSIAN MALACOLOGICAL LITERATURE remains largely
unknown in the scientific community of the Free World
until it is translated into a Western language, or cited in
the Zoological Record. The Referativnyy Zhurnal, a
publication of the Government Committee of the Soviet
Ministry of the USSR for Science and Technology (Gosu-
darstvennyy komitet soveta ministrov CCCP po nauke i
tekhnike) and the Academy of Science (Akademiya nauk
soyuza sovetskikh sotsialisticheskikh respublik), is issued
monthly in Moscow by the All Union Institute for Scientif-
ic and Technological Information (Vsesoyuznyy institut
nauchnoy i teknicheskoy informatsii). Section D in Bio-
logy considers General Zoology, Invertebrate Zoology and
Hydrobiology. The neontological literature on mollusks is
under Invertebrate Zoology.
World literature is considered, and for most papers, a
Russian translation of the author’s original abstract or a
résumé written by a Russian authority, is included.
I have reviewed the 12 issues of the Referatiunyy Zhur-
nal for the year 1968 and translated the titles of the
Russian articles contained therein. Although the volume
of literature is not as great as might have been anticipated
(paleontological papers are not included), certain book
length items and symposia are significant. The date on
all papers published in 1967 has been omitted. I have
abbreviated the following:
ZZ Zoologicheskii Zhurnal (Zoological Journal)
BWHO Byul. Vsemiri. Organiz. Zdravookhr. (Bulletin
of the World Health Organization)
TZIL Trudi zool. inst. Akad. nauk (Works of the Zoo-
logical Institute of the Academy of Science of
the USSR, Leningrad)
ES English summary
I am indebted to Mr. Vojislav Jovanovié of the Uni-
versity of Belgrade for checking the accuracy of my trans-
lations.
Auimoy, A. FE
On the possible role of animal filtrators in the process of self-puri-
fication of water bodies, on the example of the population of the
freshwater mollusc Sphaerium corneum (1). TZIL, 42: 305 - 311
Authors
Mollusks and their role in the ecosystem. Acad. Nauk SSSR, Zool.
Inst. Leningrad, Nauka, 95 pp., illus.
THE VELIGER
Vol. 12; No. 2
Barat, A. A.
Some information on the biology of the squids of the southeastern
region of the Pacific Ocean. Rybnoe Khoziaistvo. No. 8. pp. 15 - 17
BEsHEVLI, L. E. «& V. A. KoLyacin
On the discovery of the mollusk Mya arenaria (Bivalvia) in the
northwestern part of the Black Sea. Vestnik zoologii. No. 3, pp. 82
to 84
Borovyacin, V. L. « D. A. M. SakHaRov
1968. The ultrastructure of the gigantic (nerve) axons of triton.
Atlas Nauka. 100 pp., illus.
BurcHins’kKA, L. FE
1968. Histochemical peculiarities of the nerves of Planorbis cor-
neus. Fiziol. Zhur. 14 (2): 266 - 267 (Ukrainian)
Butenxko, Yu. V.
On the freshwater molluscan fauna of southern and southeastern
Kazakhstan. TZIL, 42: 205 - 212
Crip.anp, C. C.
The stability of the mollusks Bulinus (Physopsis) globosus, Bulinus
(Ph.) africanus, Biomphalaria pfeifferi and Lymnaea natalensis to
desiccation under experimental conditions. BWHO, 36 (3): 497 to
503
Dawoop, I. « B. Dazo
Field trials of two new molluscicides (molyutseda and WL 8008)
in the district of project “Egipet-49.”” BWHO, 35 (6) : 439 - 946
Go.ixov, A. N. & O. A. ScarLato
The mollusks of Possiet Bay (Japan Sea) and their ecology.
TZIL, 42: 5 - 154
GortysuHkIna, R. A.
The mollusks of the Angara River. Izv. giol. geogr. sci., Inst.
Irkutsk. Univ., 20: 65 - 94
Grossu, A. V.
Caucasian element in the malacofauna of Rumania. TZIL 42:
155 - 158
Karnaykuov, V.N., E. V. Mev’nikova, V. A. SvorEN’ & R. T. Fin
1968.Comparative microspectral investigations of the central ner-
vous systems of mollusks. Biofizika, 13 (3) : 477 - 482
Kiseeva, M. I.
Additions to the gastropod fauna of the Black Sea. ZZ 46 (5):
764 - 765 (ES)
KonpbraT Ev, G. P.
On the methods of studying the filtering rate in mollusks. (Jn)
Fauna of the reservoir of Volgograd and its influence on pollution.
Saratov, Saratovsk. Universitet, pp. 100 - 106
On the extent of mineralogical formation of bivalve mollusks in
the Volgograd reservoir. ibid. pp. 16 - 24
Kructoy, N. D.
On the biology of the little pond snails (Lymnaea) in the region
of Smolensk. ZZ 46 (5): 676-679 (ES)
Lazareva, A. I. 5
On the systematics of the small pond snails (Family Lymnaeidae,
Gastropoda Pulmonata) from Kazakhstan. TZIL 42: 198 - 204
On the systematics of the small pond snails of Kazakhstan from
the group of Lymnaea palustris MiLLER (Gastropoda, Pulmonata).
ZZ 46 (9): 1340-1349 (ES)
Lrxwarev, I. M. & YA. I. StaRoBocaTov
Materials on the mollusk fauna of Afganistan. TZIL 42: 159-197
LocvinENKO, B. M. « Ya. I. STAROBOGATOV
On the study of species composition of bivalve mollusks in
thanatocoenoses on the underwater slope of the Azerbaidzhan
coast. (In) Opyt geologo-geomorfol. i gidrobiol. issled. beregovoy
zony morya. Leningrad, Nauka. pp. 225 - 235
Vol. 12; No. 2
THE VELIGER
Page 227
Meyuine, A. H., J. Meyiinc « R. J. PircHrorp
1966. Physico-chemical features of the matters used as mollusci-
cides, hydrolyzed sodium-tritilmorfolina (Sheel WL 8008). BWHO
35 (5): 759 - 762
MIKHEEV, V. P.
Filter feeding on Dreissena. Trudi Vsesoyuznyy nauchno-issledo-
vatel’skii institut prud. Rybnogo Khoziaistva, 15: 117 - 129 (ES)
Miteykovskiy, S. A.
The larval morphologies of the Monoplacophora and bivalved
gastropods and their position in the system of molluscan types.
ZZ 46 (6): 850 - 855 (ES)
Natsviisucuviui, M. G.
On studies on the Jand mollusks of the vicinity of Tiflis. Soobsh-
cheniia Akademii Nauk Gruzinskoi SSR. 48 (1): 191 - 194
NEyYFERT, A. V.
The rate of filtration and the passage of food through the diges-
tive tract of Cardium edule (L.) (In) Data on the biocoenosis and
biology of benthic organisms in the Black Sea. Kiev, Science Memo-
rial, pp. 113 - 121
Niko acy, V. A.
On the mass reproduction of slugs in the region of Orlov.
Uchenye zapiski Orlovskogo gosudarstvennogo pedagogicheskogo in-
stituta. 35: 117 - 122
On the forecast of mass reproduction of slugs in the region of
Orlov. (In) Materials of the third zoological congress, pedagological
institute of the Russian soviet federation of socialist republics. pp.
326 - 329
Morphological and biological features of the egg of the reticulated
slug (Deroceras reticulatum Mu.x.). Uchenye zapiski Orlovskogo
gosudarstvennogo pedagogicheskogo instituta, 35: 123 - 128
Reznix, Z. V.
Investigations of the fauna of land mollusks in the Stavropol
region. Sbornik statey po zool., Krasnodar, pp. 97 - 101
Sav’skiy, V. A.
1968. On some distinctions of syndesmyas -- Abra (Syndesmya)
Puit., acclimated in the Caspian Sea from the Azov Sea. Ecological
biogeography of the contact zone of the sea. Kiev, Science Memo-
rial, pp. 91 - 95 :
Suirr, K.
1966. Experiments with the use of the chemical Sodium-tritil-
morfolina (Shell WL8008) in its role as a molluscicide in Southern
Rhodesia. BWHO, 35 (2): 205 - 214
SHILEYKO, A. A.
On variable characteristics of some land mollusks. Vestnik, Mos-
kov. Universitet. Seriia Biologii, Pochvovedeniia, no. 2, pp. 14-21
Systematic notes on the genus Helicella Fer. (Pulmonata, Heli-
cidae) with descriptions of new species from Talysha. Vestnik. Mos-
kov. Universitet. Seriia Biologii, Pochovedeniia, no. 4, pp. 27 - 34
Materials on the molluscan fauna of the family Ellobiidae in
Talysha. Nauchnye doklady Vysshei Shkoli biologicheskie nauki.
No. 8, pp. 20 - 23
On the ecology of the land mollusks of the Kola Peninsula.
Byulleten Moskovskogo obshchestva ispytatelei prirody, Otdel bio-
logii. 72 (6): 42-45 (ES)
On the reproductive biology and the adaptation of the juvenile of
the slug Parmacella ibera. ZZ 46 (6): 948 - 949 (ES)
On the question of the variability of a lymnaeid (Mollusca,
Gastropoda) in the Kola Peninsula. Gidrobiol. Zhurnal. 3 (3):
77 - 80
1968. Mollusks of the family Helicidae (Pulmonata:Stylomma-
tophora) from Talysha. ZZ, 47 (3) : 337 - 347 (ES)
Smirnova, N. F
1968. On the existence of two forms of the edible bivalve mollusk
Crenomytilus grayanus DunKer. Doklady Akad. Nauk SSSR.,
179 (3): 742-745
Soxo.ov, E. N., G. G. ArRAKELov & L. B. Levinson
Parallel excitation with spontaneously discharging neurons in
ganglia of the little pond snail, Limnaea stagnalis. (In) Evolution
neurophysiol. and neurochem., Nauka, Leningrad, pp. 3-11 (ES)
Soxo ov, V. A.
On the pedal-visceral connective in the nervous system of the
pearly mussel Unio pictorum. (In) Evolution. neurophysiol. and
neurochem., Nauka, Leningrad, pp. 26-30 (ES)
Spassxiy, N. N.
Life cycle of the mollusk Adacna vitrea and seasonal dynamics of
its biomass and numbers. Trudi Kaspiiskii nauchno-issledovate’skii
institut Khoziaistva. 23: 91 - 94
STADNICHENKO, A. P.
1968. Phospholipids in digestive glands of mollusks parasitized by
a larval trematod. Parazitologiya 2 (2): 177-178 (ES)
SrarosocarTov, Ya. I.
On the construction of a system for freshwater pulmonate mol-
lusks. TZIL 42: 280 - 304
SraropocaTov, YA. I. & E. A. STRELETSKAYA
The composition and the zoogeographic characteristics of the
freshwater mollusk fauna of eastern Siberia and the northern Far
East. TZIL 42: 221 - 268
TatisHvitt, K.G.,K.G. BaGDASARYAN & ZH.R.M.KAZAKHASHVILI
1968. Reference book on the ecology of marine gastropods. The
way of life of gastropod mollusks, belonging to the generic types
present in the Cenozoic of the South of the USSR. Nauka, 169 pp.
Turpaey, T. M., C. H. Nistratova & D. A. SAKHAROV
The evolution of cholinergic regulation of the activity of the
heart in mollusks. Zhurnal Obshaia Biol. 28: 618 - 626 (ES)
Usnakovy, B. P. « I. M. PasHKOVA
Intrapopulation variation of heat resistance of muscles and
glycerin-prepared muscle strands in different populations of Black
Sea midi [Mytilus?]. (In) The variation of heat resistance of
animal cells in ontogeny and phylogeny. Leningrad, Nauka, pp.
74-81 (ES)
Uva.ieva, K. K.
New species of land mollusks from the southern Altai. TZIL, 42:
213 - 220
Vinnikov, Ya. A., O. G. GazENnxo et al.
1968. Cyto-chemical and electron-microscopical investigations on
the statocysts of cephalopod mollusks. Zhurnal Evolyuts. biokhimini
i fiziol. 4 (1): 10-23 (ES)
Wricurt, C. A. « G. C. Ross
1966. Investigations on the egg albumins of the mollusks Bulinus
africanus and B. forskalii by an electrophoretic method. BWHO.
35 (5): 749-758
Page 228
Report on some Abnormal Chitons
from California and British Columbia
BY
GLENN BURGHARDT
California Academy of Sciences, San Francisco
California 94118
AND
LAURA BURGHARDT
14453 Nassau Road, San Leandro, California 94577
IT CAN GENERALLY BE SAID that chitons have 8 plates. But
when collecting these mollusks one finds that this is not
always the case. Specimens with 6, 7, or even 9 plates
are sometimes found. CHacE & CHAcE (1930) and RoTtH
(1966) list a total of 14 specimens collected with less than
the normal 8 valves or plates. Berry (1935) lists a
specimen with 9 valves. Bringing the records up to date,
we Offer a list of 24 specimens which have less than the
normal 8 valves. These specimens have all been collected
in California, with one exception.
Cyanoplax dentiens (Gouxp, 1846)
(1) collected May 14, 1967 on a —0.6 foot (ft. here-
after) tide at Frenchman’s Reef, San Mateo County,
California; 8.5 mm long, 5.6 mm wide in dry condi-
tion.
(2) coll. March 24, 1968, — 0.6 ft. tide at Pigeon Point,
San Mateo County, California by Mrs. John (Salle)
Crittenden; 10.0mm long and 6.0mm wide in al-
cohol (Crittenden collection).
Ischnochiton radians CARPENTER in Pitssry, 1892
collected February 2, 1969 on a — 0.7 ft. tide at Carmel,
Monterey County, California by Mike Burghardt;
22.3 mm long and 13.7 mm wide in alcohol.
Katharina tunicata (Woop, 1815)
collected June 23, 1966 on a 0.9 ft. tide at Victoria,
British Columbia, Canada; 61.6 mm long and 39.5
mm wide in dry condition.
Lepidozona californiensis (BERRY, 1931)
collected December 30, 1967 on a —1.8 ft. tide at
Doheny Beach, Dana Point, Orange County, Cali-
fornia; 4 specimens: 7.8mm X 5.0mm, 10.8mm
7.1mm, 19.1 mm 11.9 mm, and 20.6mm x 12.3
mm in length and width, respectively, in alcohol.
Lepidozona cooperi (Pirssry, 1892)
(1) collected in 1958 at “Monolith Beach” near Pes-
cadero, San Mateo County, California; 26.7 mm long
and 14.7 mm wide in semi-curled dry specimen.
(2) collected July 4, 1966 on a —0.9 ft. tide at. Moss
Beach, San Mateo County, California; 23.5mm
THE VELIGER
Vol. 12; No. 2
long and 16.0 mm wide in dry condition.
(3) collected May 18, 1968 on a —0.6 ft. tide at Bo-
dega Bay (breakwater), Sonoma County, California
by Mrs. John (Salle) Crittenden; 29.0 mm long and
18.5 mm wide in alcohol (Crittenden collection).
Mopalia ciliata (SowERBy, 1840)
(1) collected November 12, 1966 on a — 1.2 ft. tide at
Pigeon Point, San Mateo County, California; 32.0
mm long and 20.0 mm wide in dry condition.
(2) collected March 24, 1968 on a —0.6 ft. tide at
Pigeon Point, San Mateo County, California by Mrs.
John (Salle) Crittenden; 26.5 mm long and 17.8 mm
wide in alcohol (Crittenden collection).
(3) collected May 18, 1968 on a -0.6 ft. tide at Bode-
ga Bay (breakwater), Sonoma County, California;
30.7 mm long and 22.2 mm wide in alcohol.
(4) collected June 28, 1968 at 50 - 60 ft. with SCU-
BA gear by Walter Schneebeli in Monterey Bay (off
breakwater), Monterey County, California; 34.3 mm
long and 22.9 mm wide in alcohol.
(5) collected March 23, 1969 on a — 0.6 ft. tide at Bo-
dega Bay (breakwater), Sonoma County, California ;
15.3 mm long and 8.6 mm wide in alcohol.
Mopalia lignosa (Goutp. 1846)
(1) collected June 11, 1968 on a — 2.1 ft. tide at Car-
mel, Monterey County, California; 17.2 mm long and
13.9 mm wide in alcohol.
(2) collected at the same station as the preceding spe-
cimen, by Mrs. John (Salle) Crittenden; this six-
plate specimen measures 37.0 mm long and 27.9 mm
wide in alcohol (Crittenden collection).
Mopalia muscosa (Goutp, 1846)
collected March 5, 1967 on a — 0.5 ft. tide at French-
man’s Reef, San Mateo County, California; 36.6 mm
long and 23.1 mm wide in dry condition.
Nuttallina californica (NUTTALL in REEvE, 1847)
collected May 22, 1966 on a — 1.3 ft. tide at Carmel,
Monterey County, California by Mike Burghardt;
16.3 mm long and 6.7 mm wide in dry condition.
Nuttallina fluxa (CARPENTER, 1864)
collected December 27, 1966 on a —1.1 ft. tide at Re-
fugio Beach, San Luis Obispo County, California;
11.3 mm long and 8.3 mm wide in semi-curled dry
condition.
Stenoplax heathiana Berry, 1945
collected March 5, 1967 on a —0.5 ft. tide at French-
man’s Reef, San Mateo County, California; 60.1 mm
long and 24.5 mm wide in dry condition (re-articu-
lated).
Tonicella lineata (Woop, 1815)
(1) collected July 4, 1966 on a —0.9 ft. tide at Moss
Beach, San Mateo County, California; 9.0 mm long
and 6.6 mm wide in dry condition.
Vol. 12; No. 2
(2) collected March 24, 1968 on a —0.6 ft. tide at
Pigeon Point, San Mateo County, California; 22.2
mm long and 16.5 mm wide in alcohol.
Unless otherwise noted all of the specimens listed above
were collected by the authors and are in their private
collection.
LITERATURE CITED
Berry, SAMUEL STILLMAN
1935. A further record of a chiton (Nuttallina) with nine
valves. The Nautilus 48 (3): 89-90 (January 1935)
Cuacr, Emery Perkins & ELste MarcAarRET Hersst CHACE
1930. Two seven-valved chitons from Mendocino, California.
The Nautilus 44 (1): 7-8 (July 1930)
Rotn, Barry
1966. | Some abnormal chitons from Washington State. The
Veliger 9 (2): 249 - 250; 1 text fig. (1 October 1966)
New Range for
Mopalia hindsi recurvans BARNAWELL, 1960
BY
GLENN BURGHARDT
California Academy of Sciences, San Francisco
California 94118
AND
LAURA BURGHARDT
14453 Nassau Road, San Leandro, California 94577
PreviousLy Mopalia hindsi recurvans BARNAWELL, 1960
has been restricted in recorded locality to San Francisco
Bay, California. BARNAWELL states that he found the sub-
species living with M. h. hindsi (REEvE, 1847) which
outnumbered the new taxon by a ratio of 9 : 1. In recent
months, while we were collecting on the breakwater at
Doran County Park, Bodega Bay, California we obtained
a number of this subspecies. This extends the recorded
range 14 miles to the north and indicates that this sub-
species is not limited to the San Francisco Bay as previous-
ly believed. This time M. h. recurvans outnumbered M.
h. hinds: by a ratio of 4 : 1. Another interesting fact was
that the subspecies M/. h. recurvans was found living on
the channel side of the breakwater where the current was
fairly strong and the water clear while the M. h. hindsi
were generally found on the bay side of the breakwater
where the water was calmer but the animals were covered
with mud and silt. However, this was not the case in 100%
of the instances.
Specimens from the Bodega Bay colony were compared
with Paratypes nos. 2 and 3 in the California Academy of
Sciences collection (CAS nos. 12141 and 12142) and
were found to be identical. The taxonomic characteristics
THE VELIGER
Page 229
mentioned by BARNAWELL (1960) are easily noted in this
colony. With both Mopalia hindsi hindsi and M. h. re-i
curvans living in the same locality and with a strong pref-
erence for a certain ecological location it would seem that
there is a good case for raising the subspecies to specific
rank. Further study may resolve the problem.
LITERATURE CITED
BARNAWELL, EARL BAKER
1960. Mopalia hindsi recurvans, subspec. nov. (Amphineura)
The Veliger 3 (2): 37-40; plt. 6 (1 October 1960)
Range Extension of Tylodina fungina
in the Gulf of California
BY
JAMES W. McBETH
AND
R. DAVID BOWLUS
University of California at San Diego
Scripps Institution of Oceanography, La Jolla,
California 92037
On May 23, 1969, one living specimen of Tylodina fungina
Gass, 1865 was obtained while we were SCUBA diving
in 30 feet of water on the west side of Isla Espiritu Santo
in the Gulf of California, Mexico. It was found on a
sponge with a purple surface on an area where the yellow
internal tissues were exposed, thus suggesting feeding ac-
tivity by the opisthobranch. The interior tissues of the
sponge were similar in color and texture to Verongia
thiona DE LAUBENFELS, 1930, the sponge upon which
Tylodina occurs in southern California (Lance, 1961).
Tylodina fungina has been reported from the Gulf of
California only once before (DuSHaNE, 1966). In this
instance one specimen was collected intertidally at Guay-
mas, about 300 miles due north of Isla Espiritu Santo.
Tylodina has been reported along the Pacific coast from
Cayucos, California, to Todos Santos, Mexico (SPHON &
Lance, 1968).
LITERATURE CITED
DuSuan_e, HELEN
1966. Range extension for Tylodina fungina Gass, 1865 (Gast-
ropoda) . The Veliger 9 (1) 86 (1 July 1966)
Lance, JAMES ROBERT
1961. A distributional list of southern California opistho-
branchs. The Veliger 4 (2) : 64-69 (1 October 1961)
Spuon, Gate G., Jr., « JAMES RoBertT LANCE
1968. An annotated list of nudibranchs and their allies from
Santa Barbara County, California. Proc. Calif: Acad. Sci.
(4) 36 (3): 73-84; 1 fig. (25 September 1968)
Page 230
Ascophyllum nodosum:
A Source of Exotic Invertebrates
Introduced into West Coast
Near-Shore Marine Waters
BY
RICHARD L. MILLER
Department of Biology, Temple University
Philadelphia, Pennsylvania 19122
THE INTRODUCTION of marine mollusks into California
waters has been discussed recently by Cartton (1969).
He concerns himself with 2 possible methods of introduc-
tion of Littorina littorea LINNAEUS, 1758 into San Fran-
cisco Bay:
1. with shipments of Atlantic oysters, Crassostrea
virginica (GMELIN, 1791), planted in the Bay, or
2. with shipments of the Atlantic quahog, Mercena-
ria mercenaria (LINNAEUS, 1758), either planted
in the Bay or dumped into the Bay by shippers
after spoilage.
Since both the oyster and the quahog are mud dwellers
it is not unlikely that closely associated organisms would be
carried along with them, particularly if the animals were
shipped in or with substrate. This might easily explain the
introduction of mud or sand dwellers such as Nassarius
obsoletus (Say, 1882), Gemma gemma (TotTTEN, 1834),
Busycotypus canaliculatus (LINNAEUS, 1758), or Uro-
salpinx cinereus (Say, 1882), a predatory associate of
oysters. It would not account for the introduction of a
rocky intertidal inhabitant like Littorina, however.
There is another method of organism introduction into
San Francisco Bay, and probably many other bays on the
West Coast, that is not only continually on the increase
but is a source of dozens of exotic species of invertebrates
and algae. The source is Ascophyllum nodosum (LINNAE-
us) Le Jouts, 1829 (and many members of the genus
Fucus) used as packing material to protect the American
lobster Homarus americanus MiLtnr-Epwarps, 1837
during air shipment from the northeastern United States.
These fucoids characteristically bear a small population
of invertebrates and algae, most of which are inhabitants
of the upper to mid-intertidal. Since shipment is rapid,
most of these forms survive and can be introduced into
possibly suitable environments if the alga is dumped into
local marine waters within several hours after arrival. I
have obtained animals from these shipments which have
been successfully maintained at marine stations and have
THE VELIGER
Vol. 12; No. 2
sometimes used algae sent with lobsters as a source of
experimental material when short of funds.
In 1966, I talked to a shipper near Fisherman’s Wharf
who supplies many of the restaurants in the Bay area
and was told that it was his practice to dump the algae
into the Bay. Since several shipments arrive per month
and the demand for lobsters is unlikely to decrease, we can
expect that an increasing number of potential seed organ-
isms will be released into the Bay in the future. I have
found the following organisms to be relatively common
survivors in these shipments:
Coelenterata
Campanularia flexuosa (Hincxs, 1857)
Clava leptostyla Acassiz, 1862
Sertularia pumila LinnaEus, 1758
Platyhelminthes
Monoophorum sp.
Monocoelis sp.
(several other unidentified Rhabdocoels and Alleo-
coels)
Annelida
Spirorbis sp.
Crustacea
Ampelisca sp.
(several unidentified gammarid amphipods)
Mollusca
Littorina littorea LINNAEUS, 1758
Littorina obtusata Linnaeus, 1758
Mytilus edulis Linnagus, 1758
Mitrella lunata Say, 1882
Ectoprocta
Flustrellidra sp.
Bowerbankia sp.
Other species, less commonly found, are:
Porifera
Leucosolenia sp.
Coelenterata
Tubularia sp.
Obelia sp.
Gonothyrea lovent (ALLMAN, 1859)
Annelida
several species of small errant worms
Crustacea
Balanus amphitrite niveus Darwin, 1854
Echinodermata
Astertias forbesi (Desor, 1848)
Ectoprocta
Bugula sp.
The numbers and species composition in any one ship-
ment vary widely depending on the season of the year and
presumably the area from which the algae were origi-
nally collected. The resistance of these organisms to
Vol. 12; No. 2
desiccation varies considerably with the species, and the
overall prevalence of the forms listed above reflects this.
Some of the algae found include:
Chaetomorpha area Kutzine, 1845
Sphacelaria cirrosa AGARDH, 1824
Polysiphonia sp.
Ulva lactuca Linnagus, 1758
Ceramium sp.
Cladophora sp.
Enteromorpha sp.
Investigators should be aware that these organisms are
being introduced now and have been introduced in the
past in this manner. Since Ascophyllum floats, probably
the entire Bay is subject to colonization by the organisms
clinging to it. What is surprising is the lack of reports of
these organisms. This could be ascribed to the resistance
of the local fauna and flora to influxes of new forms
(especially in the intertidal, where space competition is
high), the presence of many of them as part of the already
established fauna and flora, or simply the lack of system-
atic collecting in the Bay. It would seem worthwhile for
someone to determine accurately the numbers and spe-
cies composition of the viable organisms in Ascophyllum
shipped with lobsters over a long period so that we know
just what is entering the Bay by this route. Experiments
designed to study the mode of distribution of these forms
and their rates of survival in competition with the local
fauna might be highly informative.
LITERATURE CITED
Carton, JAMES
1969. = Littorina littorea in California (San Francisco and Trin-
idad Bays). The Veliger 11 (3): 283-284 (1 Jan. 1969)
Cadlina modesta:
A Range Extension, with Notes
on Habitat and a Color Variation
BY
HANS BERTSCH
Franciscan School of Theology, 1712 Euclid Avenue
Berkeley, California 94709
(1 Text figure)
IN THE ORIGINAL DESCRIPTION of Cadlina modesta Mac-
FarLAnp, 1966 (Nudibranchia : Dorididae), the species
was recorded from Point Pinos and Point Cabrillo (Pacif-
THE VELIGER
Page 231
ic Grove near Monterey, California, 36°36’N, 121°54’W)
and from La Jolla, California (32°52’N, 117°15’W).
Since then, its occurrence at intermediate points in the
range has been established: SpHON & Lance (1968),
“frequent” occurrence in Santa Barbara County; and
Rotter & Lone (1969), “common” occurrence in San
Luis Obispo County.
On April 8, 1969 (a —0.7 foot tide), I obtained 3
specimens that were crawling on the bottom of rocky
tide pools in the middle to low intertidal zone (Ricketts
& Cavin, 1968) at Moss Beach (37°31’N, 122°31’W),
northern San Mateo County. This is an 85-mile north-
ward extension of the range (measured along the coast-
line), and a new record for San Mateo County.
These specimens were kept alive in a one-gallon tank
for about 36 hours. During this period I observed them
crawling over some shoots of the alga Gastroclonium
coultert (Harvey, 1853) at 7 different occasions. Three
other species of nudibranchs [ Tritonia festiva (STEARNS,
Figure 1
Sketch of Cadlina modesta MacFartanp, 1966
A: Specimen with complete series of lemon-yellow spots
B: Specimen without spots in front of rhinophores (gills drawn
retracted )
1873), Diaulula sandiegensis (Cooper, 1862) and Cad-
lina luteomarginata MacFartanp, 1966] from Moss
Beach were kept in another one-gallon tank, but I did
not observe them in association with G. coulteri during
this 36-hour period of random observation. Gastroclonium
coulteri was placed in the tank because of its frequency
along the sides of the pools where C. modesta was
obtained.
On April 19, 1969, I was again collecting at Moss Beach
(a —0.5 foot tide), and found 2 additional specimens of
Cadlina modesta in similar tide pool habitats. These were
crawling over the calcareous coralline alga Corallina sp.
when collected. Since MacFarLanb rather broadly de-
scribed the habitat of these animals as “rocky open pools”
Page 232
and the Large Tide Pool at Point Pinos, the association
of C. modesta with these 2 forms of algae is mentioned.
Three of the specimens collected exhibited a color vari-
ance from MacFar.anp’s original description. He states
that on the notum there is “a continuous irregular series
of small lemon-yellow spots” that “extend around in
front of the rhinophores.” These 3 animals lacked these
prominent spots in front of their rhinophores. Otherwise,
the marginal series of spots was continuous around the
notum. The other specimens closely matched MacFar-
LAND’s color description. Although the 5 specimens varied
in notal length from 16 to 33 mm and in width from 8
to 13 mm, there was not a sufficient sample to establish
any significant correlation between the animal’s size and
the absence of the lemon-yellow spots in front of the
rhinophores. The color of the rhinophores also varied from
whitish to a dusky brown. MacFarianp wrote that the
rhinophores were seldom dusky.
Mr. Steven J. Long has informed me that these color
variations occur also in animals from San Luis Obispo
County. Animals of the originally described color pattern
have been collected in that county, too.
The specimens collected April 8, 1969, are in the col-
lection of the California Academy of Sciences to permit
confirmation of the identification.
I gratefully acknowledge the help of many people in
preparing this note, especially Dr. Paul Silva for his
kindness in identifying the algae, and Mr. Allyn G. Smith
for his generous and helpful criticism.
LITERATURE CITED
MacFar.anp, Frank Mace
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
Ricketts, Epwarp F. « Jack CALvIN
1968. Between Pacific tides. 4th ed., rev. by J. W. Henc-
PETH. xiv +614 pp.; illus. Stanford Univ. Press, Stanford,
Calif.
Ro.ier, Ricuarp A. & STEVEN J. Lone
1969. An annotated list of opisthobranchs from San Luis Obis-
po County, California. The Veliger 11 (4): 424-430; 1
map (1 April 1969)
Spuon, Gate G., Jr. & JAMES RoBert LANCE
1968. An annotated list of nudibranchs and their allies from
Santa Barbara County, California. Proc. Calif. Acad.
Sci. 36 (3): 73-84; 1 fig. (25 September 1968)
fe
THE VELIGER
Vol. 12; No. 2
A Note on the Opsithobranchs
of Santa Cruz Island, California
BY
STEVEN J. LONG
126 Esparto Avenue, Pismo Beach, California 93449
On Marcu 15, 1969, the Conchological Club of South-
ern California sponsored a trip to Fraser Point (34°04’ N;
119°56’ W) on Santa Cruz Island, California. While on
this trip, the author, along with Mr. Richard A. Roller
and Mr. Hans Bertsch, collected 24 species of opistho-
branch mollusks.
SPHON & Lance (1968) reported 21 species from
Santa Cruz Island. Although not all of them were ob-
tained, the following 13 species, which have not been
previously listed from Santa Cruz Island, were collected:
Aegires albopunctatus MacFarvanp, 1905
Aldisa sanguinea (Cooper, 1862)
Ancula pacifica MacFartanp, 1905
Coryphella trilineata O’DonocHuE, 1921
Dendronotus albus MacFaruanp, 1966
Dendronotus frondosus (Ascantus, 1774)
Discodoris heathi MacFarianp, 1905
Doto amyra Marcus, 1961
Melibe leonina (Goutp, 1853)
Polycera atra MAcFaruanp, 1905
Rostanga pulchra MacFarvanp, 1905
Triopha carpentert (STEARNS, 1873)
Tritonia festiva (STEARNS, 1873)
One of these 13 species, Ancula pacifica, is also a new
record for Santa Barbara County.
LITERATURE CITED
Spuon, Gate G., Jr. & James Ropert Lance
1968. An annotated list of nudibranchs and their allies from
Santa Barbara County, California. Proc. Calif: Acad. Sci.
(4) 36 (3): 73 - 84; 1 fig. (25 September 1968)
French Association of Malacology
THIS NEW ASSOCIATION was formed on January 8, 1969,
in Paris. Its purpose is to encourage studies about mollusks
and to improve relations between malacologists. As for
the activities of the association, it is anticipated to distrib-
ute bibliographic references of its members’ publications
and to organize scientific meetings. In February the Asso-
Vol. 12; No. 2
ciation consisted of 100 members, divided into 3 groups:
biologists, ecologists, and paleontologists.
The officers of the association are:
President: AtBerT Lucas (Biologist)
Vice-President: Mme. GENEvIEVE TERMIER (Paleontol.)
Secretary: BERNARD SALvaT (Ecologist).
The Association welcomes foreign members. Annual
dues are 25 French francs. The address of the society is:
Société Frangaise de Malacologie,
55, Rue Buffon, Paris 5°, France.
W. S. M.
Tue WESTERN Society oF MALAcoLocists is scheduled
to hold its Third Annual Meeting at Stanford University,
Stanford, California, June 24 to 27, 1970.
Officers and Council Members for the year are:
President: Myra KEEN
Vice-President: EUGENE Coan
Second Vice-President: BEATRICE BURCH
Secretary: CaroL SKOGLUND
Treasurer: Barry RoTH
Councillors-at-large: TwiLa BRATCHER, and ROBERT
TALMADGE
Inquiries about the meeting and applications for mem-
bership should be sent to the secretary, Mrs. Carol
Skoglund, 3846 East Highland Avenue, Phoenix, Ari-
zona 85018.
CALIFORNIA
MALACOZOOLOGICAL SocIETY, Inc.
is a non-profit educational corporation (Articles of In-
corporation No. 463389 were filed January 6, 1964 in
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THE VELIGER
Page 233
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THE VELIGER
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BOOKS, PERIODICALS, PAMPHLETS
Segunda contribuicao ao inventario dos moluscos
marinhos do nordeste Brasileiro
by Henry Ramos MATTHEWS & ELIEZER DE CARVALHO
Rios. Arquivos da Estagao de Biologia Marinha da Uni-
versidade Federal do Ceara, vol. 7, no. 2: pp. 113-121;
December 1967.
This is the second of a series of papers recording the
marine mollusks of northeastern Brazil (the first of this
series appeared in vol. 7, no. 1, pp. 67-77, 1967). The
area considered lies between the mouths of the Parnaiba
Vol. 12; No. 2
and Sao Francisco rivers, extending north to the equator,
and east to Longitude 30° West. Included are the num-
ber of specimens, catalogue number, locality, date collec-
ted, and a bibliography.
Sixty-nine species of gastropods and 34 species of pele-
cypods are recorded in this paper. Of these, “Conus ye-
majae VAN Mot, TurscH & Kemp” and “Conus selenae
Van Mot, Turscu & Kemp” are listed as new, but are
not described.
LGH
The Littoral Marine Mollusks
of Cocos-Keeling Islands (Indian Ocean)
by VirciniA Orr Maes. Proceedings of the Academy of
Natural Sciences of Philadelphia, vol. 119, no. 4; pp.
93 - 217; plts. 1 - 26; text figs. 1-4; September 6, 1967.
This paper contains an annotated list of 504 species of
marine mollusks collected by the author and a colleague,
or reported in the literature, from Cocos-Keeling Islands.
These islands (an atoll) lie in about 12° South Latitude
and 96°50’ East Longitude in the eastern Indian Ocean.
The nearest large land areas, about 600 miles northeast,
are Java and Sumatra, from which, waterborne, the
marine molluscan fauna was probably derived.
Of 379 identified forms, 82% are widely distributed in
the Indo-Pacific, 15% occur only in the west Pacific, and
3% are endemic to the Indian Ocean. None is endemic
to the Cocos-Keeling Islands. Illustrations, or references
to illustrations, of most of the species are included. Notes
on distribution, habitat, feeding, commensalism, radulae,
and observations on the soft parts of living animals are
included for many of the species.
Anyone interested in marine mollusks of the Indo-Paci-
fic will find here, in convenient form, a wealth of infor-
mation concerning an assemblage of species, many of
which range far into adjoining regions.
LGH
Les Pectinides du Miocéne de la Guadeloupe
(Antilles Frangaises)
by DentsE Monain. Bulletins of American Paleontology,
vol. 54, no. 245; pp. 471-510; plts. 40-50; 1 table.
December 2, 1968.
Fifteen species of fossil pectens from strata of Miocene
age in Guadeloupe Island, West Indies, are discussed in
this paper and 11 are illustrated. Twelve species are pos-
itively identified, 2 are compared with described species,
and 1 is identified only as to genus. Two new taxa are
THE VELIGER
Page 235
proposed, Chlamys reynali and Chlamys (Nodipecten)
colinensis (Hopson & Harris) forma guadeloupensis.
The affinities of this assemblage are chiefly with fossil
forms of Santo Domingo (Dominican Republic) and with
Jamaica, less pronounced with those of Haiti, Anguilla,
Cuba, Costa Rica, and Panama, and decidedly less with
those of southeastern United States.
The Nodipecten group is discussed and the Recent Car-
ibbean species Chlamys (Nodipecten) nodosa (LINNAEUS)
is illustrated. An excellent index adds to the usefulness
of this paper.
LGH
A Biological Survey of Bahia de Los Angeles,
Gulf of California, Mexico. III. Benthic Mollusca.
by EucEene V. Coan. Trans. San Diego Soc. Nat. Hist., vol.
15, no. 8; pp. 107 - 132, 2 figs. 25 September 1968.
This is the type of study of which many more will be
needed before we can assess properly the distributional
patterns of mollusks in the Gulf of California. The study
is necessarily limited because of sampling methods of the
expedition on which it reports. Orangepeel grab samples
were taken in a grid pattern in April, 1962 and October,
1963. Quantitative comparisons reveal surprising dis-
crepancies, especially a marked decline in numbers of
specimens and species between April and October. One
wonders whether it was due to special circumstances of
that time and place. Only repetition in another year and
sampling in a similar way other bays of the Gulf or
with different gear can answer the questions raised by
this careful analysis. The systematic list included will
supply the basic comparative data for such future studies.
It is unfortunate that in assembling the final systematic
tabulation the printer substituted “number of species” for
“number of specimens” in one column, but the reader
should be able to construe what the author originally
wrote.
MK
A New Species of the Genus Macoma (Pelecypoda)
from West American Coastal Waters, with Comments
on Macoma calcarea (Gmelin, 1791)
by Ropert M. DuNnNILL & EucENeE V. Coan. National
Museum of Canada Natural History Papers, no. 43, pp.
1-19; figs. 1-10. 9 December, 1968.
What has been identified as Macoma calcarea now ap-
pears to be comprised of two distinct species. Macoma
Page 236
elimata DUNNILL & Coan is described, being distinguished
by presence of a flattened or slightly concave escutcheon.
The range of M. elimata is from southern Alaska to
southern California, whereas M. calcarea does not range
south of Puget Sound. Both species occur along the British
Columbia and southern Alaskan coasts. They are separable
on a number of characters, of which the escutcheon is
the most readily apparent. Comparative notes are in-
cluded on spermatozoa, synonymy, and the fossil records.
MK
Descriptions of New Species of Gastropods
from Clipperton Island
by Leo G. HERTLEIN & Epwin C. ALuison. Occ. Papers,
California Acad. Sci., no. 66; pp. 1 - 13; 13 figs. 27 June,
1968.
The authors add to the knowledge of the mollusk fauna
of Clipperton Island covered by them in previous issues of
The Veliger (vol. 3, no. 1, 1960; vol. 9, no. 2, 1966). Six
new marine species in the following genera are described:
Cyclostremiscus, Amphithalamus, Odostomia, Turbonilla
(Pyrgiscus), Colubraria, and Latirus; one new land spe-
cies (Succinea atollica) is included. The species Heliacus
infundibuliformis strigatus (HANLEY, 1863), Opeas op-
aranum (PFEIFFER, 1846) and a minute marine form
questionably referred to Omalogyra are also discussed.
The figures of the minute species are excellent pen-and-
ink renditions. There is a complete list of references to the
Clipperton Island mollusk fauna.
AGS
Rare Shells
by S. Peter Dance. Univ. Calif. Press, Berkeley. 128 pp.;
24 color plates (illustrating 50 different species). $17.00.
28 April, 1969.
This book, written in the same lucid style as its prede-
cessor by the same author (Shell Collecting: an illustrated
history) was printed in Great Britain, a fact that may
account for the relatively high price of this book. We
deplore this for the simple reason that we fear that this
will keep many prospective readers of this book from
acquiring a copy, and yet, it is worthy to be added to the
library of any serious student of mollusk shells as well as
to the library of him who just likes beautiful objects.
THE VELIGER
Vol. 12; No. 2
Among the rare shells illustrated with magnificent color
figures are some of the “classical” rare shells, such as
Conus gloriamaris, several of the fabulous Cypraea spe-
cies, etc., even though quite a number of these are no
longer quite as rare as they once were. As the author
points out in one of the introductory chapters, skin divers
have contributed largely to the finding of more specimens
of these still uncommon forms. And until almost every
other shell collector has a specimen or two of each of the
50 illustrated species, this book can be considered a very
satisfactory substitute for the actual shells themselves.
RS
The Tertiary Non-Marine Mollusca of South America
by Juan J. Paropiz. Annals Carnegie Museum, vol. 40,
pp. 1 - 242; 19 plts. Carnegie Museum, Pittsburgh, Pa.
This very careful account of the known fossil mollusks
(non-marine) from South America is a scholarly work of
great value. The author describes 1 new genus, 8 new
species and introduces 2 new names.
The various chapter headings give an idea of the over-
all scope of this work: Time units and correlations; Distri-
butional data; Systematic review; Discussion and conclu-
sions; 2 appendices; notes; a summary in Spanish; and a
very comprehensive bibliography.
It is unfortunate that we cannot inform our readers
how this work may be acquired, a work which in a very
few years will surely rank among the most important
sources of information on the fossils of the entire South
American Continent, and one that will be needed by
neontological malacologists as much as by the paleonto-
logist.
RS
Seashells of North America
(A Guide to Field Identification)
by R. Tucker Assott. 280 pp.; numerous colored illus-
trations. Plastic coated paperback $3.95; clothbound $5.95
from Golden Press, New York.
It is rather amazing how much information has been
packed in this small-sized book. The illustrations are very
good, especially when one considers the method of pro-
duction.
Vol. 12; No. 2
In this book, as in all his works of a popular nature,
Dr. Abbott continues his efforts at introducing “common
names”, a tendency which is certainly not welcomed by
many people. For example, it seems to us to be just as
difficult to learn “Mera Tellin” as to learn “Tellina mera”.
Also it seems peculiar to us that so-called common names
must be invented even for species that are already known
by a common name; we have in mind Acmaea mitra,
which along the California coast (where it does occur!)
is known as “Chinaman’s hat”, yet Dr. Abbott finds it
apparently necessary to call it “white-cap limpet’.
Aside from this aspect, which probably is not as irri-
tating to others as it is to us, the book should be quite
useful for the general collector of American sea shells.
RS
Systematics, Distribution, and Abundance of the
Epiplanktonic Squid (Cephalopoda, Decapoda) Larvae
of the California Current April 1954-March 1957
by Takasu OxuTant & JoHN A. McGowan. Bull.
Scripps Inst. Oceanogr., Univ. Calif., San Diego, vol. 14;
90 pp.; 36 text figs. (many of these maps), tables. $3.00.
publ. April 14, 1969.
This is a very scholarly analysis of an enormous amount
of material collected by the California Cooperative Fish-
eries Investigations (CalCOFI), which for a period of
10 years has obtained samples at monthly intervals from
a carefully planned network of stations.
Aside from the statistical work involved in this investi-
gation, the rearing in the laboratory of some of the early
stages of different squid species is of especial interest.
For several species excellent drawings illustrate successive
stages in the development of such species as Loligo opal-
escens, Abraliopsis felis, and others. The authors are to
be commended for the completion of this very useful piece
of work.
RS
Bursatella leachi guineensis subsp. nov.
(Gastropoda, Opisthobranchia)
from Ghana
by A. Beppincton. Proceedings of the Malacological
Society of London, vol. 38, pp. 323 - 341; 1 plate; 15 text
figures; 1969.
This paper is not only a taxonomic description of a new
taxon, but at the same time a very careful anatomical,
THE VELIGER
Page 237
histological, and embryological study of the subject, an
aplysiid. However, the taxonomic treatment is somewhat
unorthodox as there is no description (diagnosis) of the
subspecies per se, only a discussion of the differences in
this form contrasted with previously described subspecies
of the nominate taxon.
RS
The Defensive Adaptations of Lima hians
(Mollusca, Bivalvia)
by T.-H. J.Gitmour. Journal of the marine biological
Association of the United Kingdom, vol. 47, pp. 209 - 221;
5 text figures; plate I; 1967.
The author, among other observations, demonstrates
that Lima hians, when attacked, discharges a substance
which is assumed to be distasteful to the presumptive
predators as it is viscous and has an acrid odor [no doubt,
to human sense organs. Ed.]. The author also observed
that the locomotor movements leading to free swimming
are the same as those used in burrowing into a substrate;
burrowing itself is an integral part of nest building. Thus,
the question as to whether Lima is a swimmer or a nest-
builder is, in GrLMour’s words “easily resolved: the two
activities are complementary.”
Many other interesting observations are presented in
this paper, for which we urge recourse to the cited paper.
Conchiglie da Collezione
(Shells to Collect)
by Sercio ANGELETTI. Istit. Geograf. Agostini - Novara,
80 pp.; 149 color illustr. $1.50 plus postage.
We saw this extraordinarily beautiful book first at the
meetings of the European Malacological Union in Vienna
and then and there purchased two copies from the photo-
grapher, Mr. Angeletti. It was, however, not until after
we had written two letters to him and one to one of our
friends in Italy that the books did finally arrive, about 8
months after we had paid for them. While this was a most
discouraging experience, the book, nevertheless is certain-
ly worth the trouble, especially in view of the unbelievably
low price. It may be necessary to emphasize that no
pretense is made in this work to give new scientific infor-
mation, but rather unusual and highly artistic color repro-
ductions of common and uncommon shells. This book,
then, would justifiably be a welcome addition to the
Page 238
library of those who appreciate the unusual beauty of
shells. It is possible that the book (which measures 9 12
inches) could be obtained from some reliable book dealers
in Italy rather than direct from the artist. We have had
highly satisfactory experiences with the Aquario di Bo-
logna, Malacological Department, Casella Postale 745,
I-40100 Bologna, Italy.
RS
Quelques Genres de Mollusques Prosobranches Marins
Inconnus ou Peu Connus de I’Afrique Occidentale
by Wituiam ADAM & JORGEN KNuDSEN. Bulletin de
l'Institut Royal des Sciences naturelles de Belgique 44;
69 pages, 36 text figures. May 14, 1969.
The following new taxa are described:
Circulus senegalensis, C. pseudopraecedens, Teinostoma
cansadoi, Leucorhynchia bicarinata, Cynisca julhieni, Tor-
nus leloupi, T. cancellatus, T. garrawayn, T: jullient, T: afri-
canus, Macromphalina dautzenbergi, Cochliolepis daut-
zenbergi, C. jullieni, and Megalomphalus mercatonis.
A map on page 3 of this paper shows the collecting
stations which range from approximately 21° N to about
15° S Latitude. But in addition to the new taxa listed, the
authors also critically examine 15 other species belonging
to these and a few other genera.
The 29 species, new and old, are all very small; but the
figures illustrating each species are very clear and ex-
tremely helpful.
RS
THE VELIGER
Vol. 12; No. 2
A Sheller’s Directory of Clubs, Books, Periodicals,
and Dealers
edited by THomas C. Rice. 76 pp. oct., mimeographed
This pocket-size pamphlet is available from “Of Sea
and Shore, Port Gamble, Washington, 98364.” We do
not know if there is a charge for this list; perhaps, since
about half of the contents is devoted to what appear as
advertisements from various dealers, there may be no
charge.
The first few pages list shell clubs and societies from the
United States as well as many foreign countries. Four
pages are devoted to various periodicals; 22 pages are
devoted to books, arranged according to various subjects
and geographical areas covered; the remainder of the
booklet contains, as already mentioned, advertisements
from dealers, again arranged according to geographical
location.
RS
A Catalog of Dealers’ Prices for Marine Shells
by THomas C. Rice. Second edition, 76 pages, mimeo-
graphed, quarto; available at $3.00 from “Of Sea and
Shore”, Port Gamble, Washington, 98364.
This appears to be a compilation from price lists of
various dealers and is obviously an effort to present more
realistically the going prices of shells than some other
publications of a similar nature have done.
RS
Statement of Ownership, Management, etc.
of “The Veliger,’ published quarterly, on the first day of July, October, January,
and April, at Berkeley, California, as required by the Act of August 24, 1912.
Publisher: California Malacozoological Society, Inc. Editor: Rudolf Stohler. Owner:
California Malacozoological Soicety, Inc., a non-profit, educational corporation.
Bondholders, mortgagees, and security holders: none.
(signed)
R. Srou er, Editor.
THE VELIGER is open to original papers pertaining to any problem concerned
with mollusks.
This is meant to make facilities available for publication of original articles
from a wide field of endeavor. Papers dealing with anatomical, cytological, distri-
butional, ecological, histological, morphological, physiological, taxonomic, etc.,
aspects of marine, freshwater or terrestrial mollusks from any region, will be
considered. Even topics only indirectly concerned with mollusks may be acceptable.
It is the editorial policy to preserve the individualistic writing style of the
author; therefore any editorial changes in a manuscript will be submitted to the
author for his approval, before going to press.
Short articles containing descriptions of new species or other taxa will be given
preferential treatment in the speed of publication provided that arrangements
have been made by the author for depositing the holotype with a recognized
public Museum. Museum numbers of the type specimens must be included in the
manuscript. Type localities must be defined as accurately as possible, with geo-
graphical longitudes and latitudes added.
Short original papers, not exceeding 500 words, may be published in the column
“NOTES and NEWS”; in this column will also appear notices of meetings of
regional, national and international malacological organizations, such as A.M. U.,
U.M.E., W.S.M., etc., as well as news items which are deemed of interest to
our Members and subscribers in general. Articles on “METHODS and TECH-
NIQUES” will be considered for publication in another column, provided that
the information is complete and techniques and methods are capable of duplication
by anyone carefully following the description given. Such articles should be mainly
original and deal with collecting, preparing, maintaining, studying, photographing,
etc., of mollusks or other invertebrates. A third column, entitled “INFORMA-
TION DESK,” will contain articles dealing with any problem pertaining to
collecting, identifying, etc., in short, problems encountered by our readers. In
contrast to other contributions, articles in this column do not necessarily contain
new and original materials. Questions to the editor, which can be answered in this
column, are invited. The column “BOOKS, PERIODICALS, and PAMPHLETS”
will attempt to bring reviews of new publications to the attention of our readers.
Also, new timely articles may be listed by title only, if this is deemed expedient.
Manuscripts should be typed in final form on a high grade white paper, not
exceeding 814” by 11”, at least double spaced and accompanied by a clear carbon
or photo copy. A pamphlet with detailed suggestions for preparing manuscripts
intended for publication in THE VELIGER is available to authors upon request.
A self-addressed envelope, sufficiently large to accommodate the pamphlet (which
measures 514,” by 81/2’), with double first class postage, should be sent with the
request to the Editor.
EDITORIAL BOARD
Dr. Donatp P. Azzort, Professor of Biology
Hopkins Marine Station of Stanford University
Dr. Jerry DonounueE, Professor of Chemistry
University of Pennsylvania, Philadelphia, and
Research Associate in the Allan Hancock Foundation
University of Southern California, Los Angeles
Dr. J. Wyatr DuruaMm, Professor of Paleontology
University of California, Berkeley
Dr. E. W. Facer, Professor of Biology
Scripps Institution of Oceanography, La Jolla
University of California at San Diego
Dr. Cavet Hanp, Professor of Zoology and
Director, Bodega Marine Laboratory
University of California, Berkeley
Dr. G Dattas Hanna, Curator
Department of Geology
California Academy of Sciences, San Francisco
Dr. Jorn W. Hepcretu, Resident Director
Marine Science Laboratory, Oregon State University
Newport, Oregon
Dr. Leo G. HERTLEIN,
Curator of Invertebrate Paleontology
California Academy of Sciences, San Francisco
EDITOR-IN-CHIEF
Dr. Rupotr SToHLER, Research Zoologist
University of California, Berkeley
Dr. A. Myra KEEN, Professor of Paleontology and
Curator of Malacology
Stanford University, Stanford, California
Dr. Victor Loosanorr, Professor of Marine Biology
Pacific Marine Station of the University of the Pacific
Dr. Joun McGowan, Associate Professor of
Oceanography
Scripps Institution of Oceanography, La Jolla
University of California at San Diego
Dr. Frank A. Pitretxa, Professor of Zoology
University of California, Berkeley
Mr. Attyn G. Smiru, Associate Curator
Department of Invertebrate Zoology
California Academy of Sciences, San Francisco
Dr. Ratpu I. Surru, Professor of Zoology
University of California, Berkeley
Dr. Cuartes R. Stasex, Associate Professor
of Zoology
Florida State University, Tallahassee, Florida
Dr. Donatp M. Witson, Professor of Biology
Department of Biological Sciences
Stanford University, Stanford, California
ASSOCIATE EDITOR
Mrs. Jean M. Cate
Los Angeles, California
(‘Weketows off Bilailinale
LIGise Gf SGOvAIos
< 3 3 qY Fu A cee Si te
Sechoman Library
A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.
Berkeley, California
VOLUME 12 January 1, 1970 NUMBER 3
CoNTENTS
The Endodontoid Land Snail Genera Pilsbrycharopa and Paryphantopsis (Mollusca :
Pulmonata) . (3 Text sna
ALAN SOLEM .. . Dee a: at aree tera tt A230
Reproductive Cycle of the Coot Gam Mone fotenat (Say), in Long Island
Sound. (Plates 37, 38; 1 Text figure)
ANTHONY CALABRESE .. . ie OE
Three New Species of Auneseean Ghtropods fon the Kasten Pacific.
(Plates 39 and 40; 4 Text figures)
WitiiaM K. Emerson & ANTHONY D’ATTILIO . . BA Dulce ae as needy 70)
A New Species of Helminthoglypta from the Mojave Deere (Plate 41; 2
Text figures)
WALTER B. MILLER : . 275
Two New Species of the Gents Cuipee BURN & Niginae raeo ( Molluscs: lex
poda : Opisthobranchia) from New Zealand Waters. (21 Text figures)
MicHAEL C. MILLER. . rE OL es ma a 490)
Occurrence of a Rare Squid, Gagnon soll AbsertSe (Rarnily Onychoteu-
thidae) in the Indian Ocean. (Plate 42; 1 Text figure)
KING UNESIS otic a0. . 290
A Note on the Ghromostne Number and Tmeerrelaionenine in ahe Marine Gastonod
Genus Thais of the United States Pacific Coast. (Plate 43)
Muzammit AHMED & ALBERT K. SPARKS. . an ELMS TAL AEDES fo a aR ets Rb BES 10 3
Five New Species of Terebra from the Eastern Poe (Plate 44)
Twita BRATCHER& R.D.BurcH . . . 205
Observations on the Anatomy and Biology of sve Galroans Wermeed Gunonots:
(Plate 45; 4 Text figures)
MicHaEL G. HapFIELD . . . Se Wiha ones Ute enim wal ce hh see OT
New Species of Panamic Marine @acropods. (Plate 46)
AME SE em VICE ENNIO Me Mica a. 1Gs7 See de (een whens! ees ge ee STO
[Continued on Inside Front Cover] Q ees 0 Na
Distributed free to Members of the California Malacozoological Society, Inc. { JUL 15 1971 \
Subscriptions (by Volume only) payable in advance to Calif: Malacozool. Soc., Inc. \
Volume 13: $18.- Domestic; $19.- in the Americas; $19.50 in all other Foreign Countries \ Ligp ARIE
Single copies this issue $14.-. Postage extra.
Send subscription orders to Mrs. JEAN M. CaTE, 12719 San Vicente Boulevard,
Los Angeles, California 90049. Address all other correspondence to Dr. R. Stouter, Editor,
Department of Zoology, University of California, Berkeley, California 94720
Second Class Postage paid at Berkeley, California
ConTENTs — Continued
Correlation of Radula Tooth Structure and Food Habits of Three Vermivorous
Species of Conus. (Plate 47)
James NyBAKKEN . . . 316
Observations on the Renroducnve Tela of he Kellet’s Whelk, Kelletia etter
(Gastropoda : Neptuneidae) . (Plates 48 and 49; 4 Text figures)
RicHAaRD J. ROSENTHAL. . - 319
The West American Species of Migergat (Gastronoda Muricidaeye including ‘Two
New Species. (Plate 50)
Emity H. VokEs . . 2) eee
Two New Epitoniidae from ne Galapagos lands ( Welbress & Greenday
(Plate 51)
HELEN DUSHANE. . . - 330
An Ecological Study of valle ores. Guopedsn in a Mixed Niesophveie Sineior
of Northern Kentucky. (7 Text figures; 3 Tables)
BRANLEY A. Branson & Donatp L, BATcH . . . 5 FAS
A New Species of Muricopsis from West Mexico. ‘(Plate 523; 4 1 Text Aeutes)h
GeorcE E. RapbwIN & ANTHONY D’ATTILIO.. . . 9 - 351
Two New Species of Deepwater Bivalves from the Gane Sen, if Plate oo
2 Text figures)
Harotp E. Vokes . . - 357
Descriptions of a New Genus and Biehe New Species af ‘Basten pecine Pierenicee
with Notes on Other Species. (Plate 54; 1 Text figure)
James H. McLean . . . . 362
On CERNOHORSKy’S Dedenation af a Lectotype fon Mu urex eamionciella LaSAIOS.
Harotp E. VoKEsS .. . . 368
A List of Recommended Nomenclannal Chances for SMnGPARL AND “Studies ee
Opisthobranchiate Mollusks of the Pacific Coast of North America.
RicHarp A:/ROLUER? = doe ho G54) a ne eo eee
NOTES. & NEWS) coos ce Re ee Te OM NE Favre 72 Sl SR So A
Range Extensions for Acanthodoris hudsoni MacFaRLAND, 1905, andOnchi-
doris bilamellata (LINNAEUS, 1767). Gary R. McDona.p
Spawning Notes V. — Acanthina angelica 1. OLDRoypD, 1918 and Acan-
thina lugubris (SowERBY, 1821). (6 Text figures)
Fay Henry WoLFson
METHODS & TECHNIQUES 2.079555 2 2°08) RRs eee ieee)
A Mechanical Shell Washer. (2 Text figures) JAcQuELin N.
MILLerR & Cart L. Hupss
BOOKS, PERTODICALS & PAMPHEEMS 2 cyte etree On
Note: The various taxa above species are indicated by the use of different type styles
as shown by the following examples, and by increasing indentation.
ORDER, Suborder, DIVISION, Subdivision, SECTION,
SUPERFAMILY, Famity, Subfamily, Genus, (Subgenus)
| New Taxa
Vol. 12; No. 3
THE VELIGER
Page 239
The Endodontid Land Snail Genera
Pilsbrycharopa and Paryphantopsis
(Mollusca : Pulmonata )
ALAN SOLEM
Field Museum of Natural History, Chicago, Illinois 60605
(3 Text figures; 6 Tables)
PUBLICATION OF A SYNOPTIC REVIEW of New Guinea and
Indonesian endodontid land snails (SoLEm, 1958) was
based upon limited material and use of only conchological
characters. Several species names were left as “Problem-
atica.” Subsequently I have been able to study nearly all
type specimens, located some previously unstudied concho-
logical material, and obtained preserved material of two
undescribed species, one belonging to Pilsbrycharopa and
the other to Paryphantopsis. A review of these two genera
is presented below in order to correct previous errors,
distinguish these genera from the endodontid land snails of
Micronesia and Polynesia, and to record anatomical data
having critical importance in interpreting pulmonate
phylogeny.
This study is a side project to monographic reviews of
the Pacific Island endodontid land snails that are com-
pleted, but whose publication will be delayed. Establish-
ment of which are systematically significant and which
are trivial characters, definition of measurements, patterns
of formal description, determination of normal variation,
plus criteria for supraspecific classification are developed
in the longer papers.
MATERIAL STUDIED
With two exceptions, specimens examined consist of type
material or specimens previously recorded in the litera-
ture. Several extremely interesting unstudied sets collected
by Lamberto Loria in Eastern Papua between 1890 and
1893 were located in the Museo Civico di Storia Naturale,
“Giacomo Doria,” Genova. Preserved specimens of two
undescribed species collected in the Eastern Highlands of
New Guinea by J. Linsley Gressitt in 1955 were found in
the Bernice P. Bishop Museum. These sets have turned a
routine conchological review of types into a study contrib-
uting data towards the problem of relating slugs to shelled
snails.
For convenience, the following set of abbreviations is
utilized throughout the text to indicate the source of
specimens examined.
ANSP Academy of Natural Sciences, Philadelphia
BMNH British Museum (Natural History), London
BPBM Bernice P. Bishop Museum, Honolulu
FMNH Field Museum of Natural History, Chicago
MHNG Museo Civico di Storia Naturale “Giacomo
Doria,” Genova
RNHL ~ Rijksmuseum van Natuurlijke Historie,
Leiden
SMF Natur-Museum Senckenberg, Frankfurt
UMMZ University of Michigan Museum of Zoology,
Ann Arbor
ZMA Zoologisch Museum, Amsterdam
ZMB Zoologisches Museum der Humboldt Uni-
versitat, Berlin
For permission to study types at their museums and for
the often very protracted loan of specimens for detailed
analysis, I am deeply indebted to Drs. R. Tucker Abbott,
Yoshio Kondo, Enrico Tortonese, C. O. van Regteren-
Altena, Adolf Zilch, Henry Coomans, Siegfried Jaeckel,
and to Mrs. W.S. S. van Benthem Jutting van der Feen,
Mr. Norman Tebble and Mr. John Peake.
Preparation of this report has been greatly aided by the
assistance of Mrs. Lynda Hanke and Mrs. Sandra Rendel-
man with slide preparation, statistical analysis of data,
and library work. Anatomical drawings in Figure 2 are
by Miss Margaret A. Moran, shell drawings in Figure 1
by Mr. Sander Nat Heilig, and Figure 3 by Mrs. Jane
Page 240
THE VELIGER
Vol. 12; No. 3
Calvin. This work was made possible by National Science
Foundation grants G-16419, GB-3347 and GB-6779.
All measurements were taken by the author. Species
less than 5 mm in diameter were measured with an ocular
micrometer under 16x - 32 magnification. Larger spe-
cies were measured to the nearest 0.1 mm with a dial
vernier caliper. In Tables 2, 3, and 4, where more than
single specimens were available, measurements are record-
ed as mean, standard error of the mean, and range of
variation, for example, “6.76++0.145 (6.27-7.32)”. Height,
diameter, and umbilical width measurements are given in
millimeters. Ribs are counted on the body whorl and
ribs/mm is an index of rib spacing on the body whorl.
Whorls are counted to the nearest eighth, “+” or “—”
indicating slight differences. Unless specified, all measure-
ments were made only on adult shells. Despite having non-
determinate growth, adult and gerontic specimens of endo-
dontids can be recognized from the shell. Measurements
of large samples collected on Pacific Islands have shown
that adult size variation shows a normal distribution
(unpublished data). Thus statistical comparisons of
samples are meaningful and useful.
DISTRIBUTIONS anp LOCALITIES
Pilsbrycharopa, as here delimited, ranges from northern
Borneo and Bali to New Britain in the Bismarck Archi-
pelago. The few and scattered locality records probably
indicate lack of collecting. Undoubtedly additional species
will be discovered. Paryphantopsis is known from the east-
em two-thirds of New Guinea and the Louisiade Archi-
pelago. Data concerning the geographic position of type
localities for the Indonesian and Bismarck Archipelago
species are provided in the original descriptions or cita-
tions, but the New Guinea localities are extremely difficult
to find on maps. Many species came from classic ornitho-
logical collecting camps, whose names are meaningless to
malacologists or cartographers. Other shells were taken
on expeditions whose itineraries would be familiar to bot-
anists or entomologists, but are unknown to scientists in
other disciplines. With the help of Dr. Austin L. Rand,
Chief Curator of Zoology at Field Museum of Natural
History and Mrs. van der Feen, I am able to provide
at least approximate coordinates for the New Guinea
localities. Accuracy of citations varies widely. Data on
the Archbold localities (see ARcHBoLD, RAND & Brass,
1942) and other West Irian localities are relatively pre-
cise. Data on the New Guinea and Papua stations some-
times are only area designations, since the original local-
ities were not precisely defined or the collections were
made long before accurate maps were available or in
uncharted areas lacking convenient reference points.
Although coordinates are cited in the text, a brief
indication of the more difficult and generalized localities
seems desirable. They are presented in rough geographic
order from west to east:
Cyclops Mountains, near the former Hollandia (= Ko-
tabaru, Sukarnapura), West Irian at about 140°35’ E,
2°30’ S;
Maeanderberg, area past great bend of Upper Sepik
River, Sepik District, New Guinea at about 141°25/
E, 4°10’ S, probably accurate to 40’;
Daulo Pass, near Chuave, Eastern Highlands, New
Guinea at about 145°10’ E, 6°03’S, accurate for
general area only;
Constantinhafen, Astrolabe Bay, Madang District, New
Guinea at about 145°45’ E, 5°20’S, accurate to
about 10’;
Vikaiku, village on Angabunga or St. Joseph River, in-
land from Hall Sound, Central District, Papua at
about 146°55’ E, 8°45’ S, visited by Lamberto Loria
in the 1890’s, almost certainly no longer in use as a
village name and thus unidentifiable;
Arva (= Aroa) River, northwest of Port Moresby,
Central District, Papua at about 146°55’ E, 9°00’ S,
exact locality along river unknown;
Moroka, near headwaters of Laloki River, east of Port
Moresby, Central District, Papua at about 147°25’
E, 9°30’ S, also a probably vanished Loria station;
Bujakori, village along Kemp Welch River, northeast of
Rigo, Central District, Papua at about 147°40’ E,
9°35’ S, another Loria station whose name may no
longer be used;
Sattelberg, Huon Peninsula, near Finschhafen, Morobe
District, New Guinea at about 147°40’ E, 6°30’ S.
Pilsbrycharopa Sotem, 1958
Archiv fiir Molluskenkunde, 87 (1-2): 24
Very small to quite large Charopinae in which the apical
sculpture consists of radial and spiral elements with vary-
ing dominance. Whorls loosely to very tightly coiled, gen-
erally about 32 to 44, more in tightly coiled species
(PR baliana, P. renschi). Spire flat to strongly elevated,
umbilicus widely open to closed. Radial sculpture normal
in most, reduced in P. gressitti and P. schneideri. Secondary
spiral cording present or absent. Whorl contour and shape
Vol. 12; No. 3
variable. Anatomy known only for one species. Pallial
cavity typically charopinine except for partial downward
rotation. Terminal genitalia distinguished by long penial
retractor muscle, elongated penis with large stimulator
but no vergic papilla, smooth and irregular pilasters, and
short epiphallus with small diverticulum.
Type species: — Pilsbrycharopa papuana SoLeM, 1958 by
OD.
Pilsbrycharopa is used in a greatly expanded sense as an
“umbrella” for several species from Indonesia, New Gui-
nea, and New Britain. Only one of the 9 species has been
dissected. Most taxa are known only from single collec-
tions of a few empty shells. Possibly 5 groups of species
should be recognized:
a) typically sculptured, flat spired or slightly elevated,
reddish brown shells with moderately to widely open um-
bilici, variable in size, apical sculpture variable.
Pilsbrycharopa kobelti (BOTTcER, 1908) from In-
donesia and New Guinea
Pilsbrycharopa timorensis (B. RENScH, 1935) from
Timor
Pilsbrycharopa brunnescens (MOLLENDORFF, 1892)
_ from Timor Laut and Papua
Pilsbrycharopa nigrofusca (SmirH, 1896) from
New Guinea (+P papuana Sotem, 1958)
b) typically sculptured, slightly elevated spire, thick
body whorl with strong lateral flattening above periphery,
reddish brown shell with narrow umbilicus, medium size,
apical sculpture of fine radials and spirals.
Pilsbrycharopa densecostulata (THIELE, 1928) from
New Guinea
c) reduced radial sculpture near aperture, very slightly
elevated spire, rounded body whorl, large flammulated
shell, umbilicus quite narrow, apical sculpture with narrow
radials more prominent than spirals.
Pilsbrycharopa gressitti, new species from New
Guinea
d) very fine radial sculpture, elevated spire of many
whorls, tight coiling pattern, laterally compressed body
whorl, reddish-yellow horn, closed umbilicus, apical sculp-
ture of radials that are lower than the spirals.
Pilsbrycharopa baliana (B. Renscu, 1930) from
Bali
Pilsbrycharopa renschi (Franc, 1952) from Flores
and Sumbawa (== Charopa vicina B. RENScH,
1930 not Preston, 1907)
e) low and broad radial sculpture with finer spirals,
strongly elevated spire, rounded body whorl, flammulated,
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Page 241
large shell, with umbilicus nearly closed by reflection and
angulation of baso-columellar margin.
Pilsbrycharopa schneideri (1. RENscH, 1937) from
Papua and New Britain
The Charopinae are widely distributed in Melanesia,
Micronesia and Polynesia as far east as the Society Is-
lands. Some 86 species have been reviewed in a just com-
pleted monograph. They are highly conservative in shell
form, basic sculptural pattern, umbilical size, whorl count
and apical sculpture. What would seem to be relatively
minor changes in the above shell features were found to
be correlated with anatomical divergences and indicated
generic separation. Besides the Pacific Islands, Charopinae
or their derivatives are major constituents of the New
Caledonian, Tasmanian and New Zealand endodontid
faunas. In Indonesia and New Guinea this group seems to
have been replaced by the various helicarionid taxa. I
have no doubt that the species in Pilsbrycharopa are
relicts. Quite possibly anatomical studies may demonstrate
that the above species belong to 4 or 5 genera. Certainly
the varying shell features listed above for groups c, d
and e were found to indicate generic differences in the
Pacific Island taxa. Since only one species could be dis-
sected and only the most fragmentary material is known
for almost all species, I prefer to define Pilsbrycharopa
in a much broader sense than genera used in the Pacific
Island monograph. The probabilities of obtaining pre-
served soft parts are remote. Division into several genera
based upon analogous variation would not provide any
phyletic insights.
Retention of a single generic name serves two practical
purposes. The species, despite their variability, do have a
basic similarity in form and sculpture that contrasts with
the form and sculpture of the New Guinea endemic genus
Paryphantopsis. The variation in apical sculpture seen in
the species lumped as Pilsbrycharopa shows how the pitted
apical sculpture of Paryphantopsis could evolve by a series
of gradual changes from the primitive charopinine pat-
tern of prominent spiral cords. This is discussed more fully
later. Classification into the two genera is practical both
from the standpoint of facilitating identification of the
two general phenotypes and in leaving open the currently
unanswerable question of which stock was the most prob-
able ancestor of Paryphantopsis. While origin of the latter
from some group of Pilsbrycharopa is reasonably certain,
without anatomical data on more species in both genera,
phyletic speculation would be of little value.
Determination of the relationships between the species
groups outlined above must await dissection of the soft
parts, but some statements can be made concerning their
relative degree of specialization. Detailed study of the
Page 242
many Polynesian-Melanesian Charopinae has included
establishing the median condition for various characters
whose states can be coded or measured. By comparing the
character states in the species groups of Pilsbrycharopa
with the median character states for the Polynesian-Mela-
nesian taxa, degrees of departure from the typical condi-
tion become obvious. Those species showing more and
greater departures from the norm are considered more
specialized than those showing greater similarity to the
median states. Such a crude comparison does not permit
relating species groups to each other, but does permit
ranking in terms of specialization.
Data on median states and quartile ranges for the tooth-
less Charopinae of Polynesia and part of Melanesia are
given in Table 1. This involves 50 species level taxa.
THE VELIGER
Vol. 12; No. 3
widely umbilicated, while P timorensis differs mainly by
its very wide umbilical opening. ‘The above 3 species have
typical apical sculpture with only slight radial elements.
Pilsbrycharopa nigrofusca is greatly enlarged, slightly
altered in proportions, and has stronger radial elements
in the apical sculpture. Except for size alterations, the
above species show few departures from the typical pat-
tern. Group “a” is thus very generalized.
Pilsbrycharopa densecostulata shows several significant
changes. The apical sculpture has narrow radial and spiral
elements, the umbilicus is much narrower, the whorl
count reduced, the H/D ratio and body whorl width
noticeably increased without prominent spire elevation,
there is strong lateral flattening of the body whorl and
apparently the ribbing is very crowded. These specializa-
Table 1
Mean Species Measurements
in Pacific Island Toothless Charopinae
Shell height in mm
Shell diameter in mm
H/D ratio
Whorl count
Diameter/umbilical width ratio
Ribs on body whorl!
Ribs/mm on body whorl!
Apical cords
irst Quartile
' Four species have reduced ribbing and were omitted from this
table
The entire 2": and 3" quartile ranges for characters are
not very large, emphasizing the relatively conservative na-
ture of this group. Apical sculpture in these species con-
sists of prominent spiral cords, sometimes reduced, rarely
with addition of secondary radial elements. Either closure
or great widening of the umbilicus is rare. Reduction of
the radial ribbing has occurred a few times and a change
to widely spaced ribbing is more common. Comparative
data on Pilsbrycharopa species are given in Tables 2 and 3.
Despite having a relatively large size, mean diameter
of the types is 3.96 mm, Pilsbrycharopa brunnescens con-
forms most closely to the median image, agreeing in H/D
ratio, whorls, D/U ratio, apertural inclination, and rib
count. The lower Ribs/mm is probably a simple correlative
of greater size. Pilsbrycharopa kobelti is small and rather
tions are not necessarily correlated with each other, al-
though the whorl thickening and compression may be
responsible for the umbilical narrowing. Hence the sepa-
ration of P densecostulata as a group “b.” Its characters
can be derived from those shown in group “a,” but I am
not claiming linear descent on the present limited evi-
dence.
Pilsbrycharopa gressitti has a flat spire, radial apical
sculpture much more prominent than the spiral, a quite
narrow umbilicus, major radial sculpture becoming very
crowded on the body whorl, retains a flammulated color
pattern, is extremely large and has a very loose coiling
pattern. Many of the same features are mentioned for
P. densecostulata, but the degree of specialization is greater
in P. gressitti, and the basic shape changes are quite dif
Vol. 12; No. 3
ferent in the two species. Pilsbrycharopa gressitti, the
only member of group “c,” shows more and stronger
specializations than does ites group “a” or “b.”
Pilsbrycharopa renschi and P. baliana have great spire
protrusion (about 0.40 body whorl width), very thick
and laterally flattened body whorls, markedly increased
whorl count, high H/D ratio, closed or nearly closed um-
bilicus, low apical radial ribs and more prominent spirals,
and very crowded radial ribbing. The specializations are
extreme in nature and group “d” is quite different from
the preceding groups.
Pilsbrycharopa schneideri has achieved many of the
group “d” changes, but in a different way. Its body whorl
is rounded and the spire very strongly protruded, its
radial apical sculpture is broadly rounded and much
more prominent than the spiral, the broad major radial
ribs are crowded and irregular on the body whorl, the
umbilicus is closed by contraction and has a peculiar re-
flection and angling of the baso-columellar lip that is
very different from the pattern seen in other Pilsbry-
charopa. The shell is very thick. Although the effect of
the changes plus the raw measurements and proportions
suggest that groups “d” and “e” are quite similar, detailed
comparison of the features fee that the same results
have been arrived at by different methods. Pilsbrycharopa
renschi and P. schneideri are specialized to a greater
degree than other species, but their similarities seem to
be convergent and there are major differences in whorl
contour, sculpture and form.
The five groups of Pilsbrycharopa show successively
greater departures from the median pattern seen in a
related and generalized genus of wide distribution with
50 known species level taxa. Group “a” is still generalized
and group “b” only moderately changed. Groups “c,” “d,”
and “e” are much more specialized, but each in a different
way or have achieved the same result by different means.
I reject any implication of a linear phyletic relationship
between groups “b-e.” Probably each group was inde-
pendently derived from ancestors sharing the character-
istics of group “a.”
In eatin: gnacies are found as far west as Borneo
(Pilsbrycharopa kobelti) and as far east as New Britain
(P schneideri), but most are from the Wallacea transi-
tion zone between the dominance of Oriental and Austral-
ian faunas (P kobelti, P timorensis, PR. brunnescens, P. bali-
ana, P. renschi) and New Guinea (P kobelti, P. brunnes-
cens, P- nigrofusca, P. densecostulata, P. gressitti and P
Schneideri). It should be noted that even with very limited
material, several species, P kobelti, P brunnescens, and P
schneideri, have wide distributions. Since land mollusks do
not show an Oriental-Australian faunal division from In-
donesia to New Guinea, this is not an important feature.
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Page 243
An artificial key to the species follows.
KEY To THE SPECIES oF Pilsbrycharopa
1. Umbilicus contained more than 15 times in the dia-
meter or closed Af cheer? Ot tee MO te ee eh ae
Umbilicus widely open or contained less than 15 times
in the diameter CP cn! on aor Prien ese ate
2. Diameter less than 4.5mm; apical sculpture fine;
crowded, narrow radial ribs sicciaitin Lathe WINS
Diameter more than 5.0 mm; apical sculpture of wide
radials; radial ribs irregular on the body whorl.
Pilsbrycharopa schneideri (I. Renscu, 1937)
3. Umbilicus closed; over 200 ribs on body whorl
Pilsbrycharopa renschi (Franc, 1952)
_Umbilicus a narrow crack; about 150 ribs on the body
whorl Pilsbrycharopa baliana (B. Renscu, 1930)
4. Adult diameter over 6.0mm Sa en ae)
Adult diameter less than 4.5 mm RS amen (
5. Umbilicus very narrow, D/U ratio more than 8; radial
Dee crowded near aperture
Pilsbrycharopa ESE, mo nov.
Umbilicus widely open, D/U ratio less than 4.50; radial
ribs not crowded near aperture
Pilsbrycharopa pienaftsa (Sus, 1896)
6. Whorls strongly flattened laterally; body whorl thick;
umbilicus relatively narrow :
Pilsbrycharopa Beiecwostetlatn (am, 1928)
Whorls not strongly flattened laterally; body whorl nar-
rower; umbilicus widely open Bends am co Se aad)
7. Diameter more than 3.0 mm ee ee ea UE
Diameter less than 2.7 mm ,
Pilsbrycharopa lala (Emam, 1908)
8. Umbilicus very widely open, D/U ratio about 2.20
Pilsbrycharopa timorensis (B. REnNscuH, 1935)
Umbilicus narrower, D/U ratio about 4.00
Pilsbrycharopa brunnescens (MG6LLENDoRFF, 1892)
Pilsbrycharopa kobelti (BOTTcER, 1908)
Charopa kobelti BortcER, 1908, Nachr. Bl. dtsch.
Malak. Gesell., 40 (4): 181-182; figs. 1-3 — Kap
Tial, Hitu, North Ambon, Indonesia.
? Charopa novoguineensis Sods, 1911, Annales Musei
Nat. Hungarici, 9: 352; fig. 7 — on trees under
Page 244
moss, Sattelberg, German New Guinea at 800 m
elevation.
Pilsbrycharopa kobelti (BOTTGER), SoLEM, 1964, Sa-
bah Soc. Journ. 2 (1-2): 23; fig. III, 23 — Go-
mantong Hill, Sabah (= North Borneo).
Beilania kobelti (BOTTGER), VAN BENTHEM JUTTING,
1964, Nova Guinea, Zool., 26: 13 - Waima, Misool.
Diagnosis: Shell very small, diameter 1.69-2.45 mm
(mean 2.06 mm), with 34 to 44 normally coiled whorls.
Apex and spire slightly to moderately and evenly elevated,
body whorl descending at most slightly more rapidly, H/D
ratio 0.442 - 0.549 (mean 0.490). Apical whorls 14, sculp-
ture of 9-12 (mean 10.8) narrow spiral cords with an
intrusion of radial swellings on last quarter to third. Post-
nuclear whorls with high, prominent, narrow, lamellar,
protractively sinuated radial ribs, 57 - 94 (mean 74.0) on
the body whorl, whose interstices are 2 to 5 times their
width. Ribs/mm 9.60 - 14.53 (mean 11.71). Microsculp-
ture of fine radial riblets, 5 to 9 between each pair of
major ribs, crossed by much finer and more crowded
spiral riblets, with relatively prominent, widely spaced
secondary spiral cording. Sutures deep, whorls strongly
rounded above, only slightly compressed laterally above
periphery and on basal margin. Umbilicus open, “U”-
shaped, slightly and regularly decoiling, contained 2.90 -
3.59 times (mean 3.17) in the diameter, margins rounded.
Aperture subcircular, very slightly compressed laterally
above periphery, inclined about 15° from shell axis.
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Vol. 12; No. 3
Pilsbrycharopa kobelti is characterized by its very small
size, nearly circular whorl contour, widely open umbilicus
and dominant apical sculpture of spiral cords with second-
ary radial elements at anterior end. Other Pilsbrycharopa
are much, much larger and have the apical sculpture
composed of more nearly equal radial and spiral elements.
Description: Shell very small, with a little less than 4
normally coiled whorls. Apex and spire slightly and evenly
elevated, body whorl not descending more rapidly, H/D
ratio 0.500. Apical whorls 14, sculpture of 9 narrow, in-
conspicuous spiral ribs with a slight intrusion of broad
radial swellings on last quarter whorl. Postnuclear whorls
with narrow, lamellar, protractively sinuated radial ribs
whose interstices are 3 to 4 times their width. Microsculp-
ture of fine radial riblets crossed by much finer and more
crowded spiral riblets with a secondary sculpture of mod-
erately prominent spiral cords. Sutures deep, whorls
strongly rounded above, not conspicuously flattened on the
outer margins. Umbilicus moderately open, “U”-shaped,
slightly and regularly decoiling, contained 3.05 times in
the diameter. Color mainly leached from the shell. Lip
and portions of body whorl badly broken. Height of holo-
type 0.95 mm, diameter 1.94 mm.
‘Holotype: Moluccas: Ambon, Kap Tial, Hitu. Natur-
Museum Senckenberg, Frankfurt number 75499.
Range: Sabah (= North Borneo), Ambon, Misool and
New Guinea.
Table 2
Variation in Group “A” Species of Pilsbrycharopa
P. kobelti (BOTtTcER)
Gomantong, Sabah
(juvenile shell)
Ambon (holotype)
Ambon (ZMA)
Waima, Misool (ZMA)
P. timorensis (B. RENscH)
holotype
P. brunnescens (M6LLENDORFF)
__Tenimber, SMF 165692-4
Vikaiku, Papua
P. nigrofusca (SMiTH)
Constantinhafen
7.19+0.577
(6.67-7.77)
7.74 £0.196
(6.90-8.42)
12.10+0.472 | 1.02+0.047
(10.46-14.53) | (0.77-1.18)
10.46 + 0.390
9.60-11.35 0.92-1.13)
not available 2.01
(2.14-2.27)
1.94+0.050
6.16+0.168 2.97 +0.045
(5.34-6.69) | (2.81-3.07)
Vol. 12; No. 3
THE VELIGER
Page 245
Material: Sabah: Gomantong Hill (2 specimens, FANH
118926). Ambon (8 specimens, ZMA collected October
11-13, 1949 by M.A. Lieftinck) : Kap Tial, Hitu (1 spe-
cimen, SMF 75499). Misool: Waima (6 specimens, ZMA
collected September 13-14, 1948 by M. A. Lieftinck).
Remarks: A complex of species may be lumped under this
specific name, but available data are insufficient to justify
any separation. The types and only known specimens of
Charopa novoguineensis were destroyed in 1956. In size |
and general appearance this shell seems to fall within the
range of variation cited above, although the brief descrip-
tion does not give sufficient details to enable placement.
There are discrepancies between the description and fig-
ures, since the description cited 44 whorls, but the figure
shows 34 whorls. I prefer to consider this a probable
synonym of Pilsbrycharopa kobelti.
Examples of “Beilania” demani (TAPPARONE-CANEFRI,
1883) have been collected on Ambon and Misool at the
same localities, and the species have been confused in
previous literature. These are easily separated by B. de-
mani having 2 prominent parietal lamellae, stronger ra-
dial ribbing, a flat spire, usually wider umbilicus and the
apical sculpture lacks the secondary radial elements.
Measurements of the several sets are given in Table 2.
The Gomantong shell is juvenile and one subadult example
is included in the measurements of the Ambon series.
An accurate rib count on the holotype was not possible and
this specimen was not directly compared with other spe-
Table 2
Variation in Group “A” Species of Pilsbrycharopa
cimens. Obviously it is within the range of variation shown
by the other specimens in respect to measurable charac-
ters. The Misool shells are smaller, slightly higher, more
narrowly umbilicated and with fewer, less crowded radial
ribs than the Ambon population. Possibly the former are
subadult, but characters indicating adulthood are poorly
defined in this species. Their lower whorl count, mean
344+ compared with 3}-+, is suggestive of their not being
adult. Without study of additional material and dissection
of the soft anatomy, I prefer to leave these as a single
rather broadly defined species.
Classification of Charopa kobelti in Pilsbrycharopa ra-
ther than one of the Melanesian-Polynesian genera is
based upon the intrusion of radial elements into the apical
sculpture, nearly circular aperture and lack of whorl
flattening, low mean whorl count and coiling pattern. The
primary alternative is an essentially Polynesian genus that
extends westward into the New Hebrides, Bismarcks and
Solomon Islands. Species of this as yet unpublished taxon
from the Bismarcks and Solomon Islands have much
tighter coiling patterns, marked lateral flattening of the
body whorl above the periphery, and no trace of radial
elements in the apical sculpture. Dissection of P kobelti
will be required in order to ascertain its generic position.
Pilsbrycharopa and the other genus differ greatly in geni-
talia so that solution of the problem will be relatively
simple. The conchological features listed above show
greater similarity to Pilsbrycharopa, hence I have classi-
fo) [}
be ‘2 3 Cr |
3 ce g By
§ Q 4 >
= — & =e
(a) qc =) QA
1.45 ! 43 3.38
1.91 oD |e 0.63 3.05
2.14+0.082 0.475 0.0071 0.69+0.032 3.11+0.042
(1.71-2.43) | (0.442-0.496) (0.51-0.79) | (2.95-3.35)
1.92+0.105 0.519+0.0110 a 0.58 +0.058 3.33 +0.160
(1.68-2.19) (0.496-0.549) = 3 0.51-0.76 2.90-3.59)
3.96 + 0.058 0.92+0.040 | 4.31+0.130
3.82-4.08) | (0.546-0.568 (0.86-1.02) | _(4.21-4.57)
3.63 + 0.056 0.89+0.021 | 4.08+0.096
3,39-3.85 0.500-0.578 35 3.79-4.56
6.43+0.112 | 0.462+0.0069 Agu 187+0.051 | 3.450.059
(6.14-6.93) | (0.439-0.489) (44-44) (1.63-2.03) | (3.31-3.76)
Page 246
THE VELIGER
Vol. 12; No. 3
fied it in that genus despite the very small size, which
corresponds more with the Bismarck-Solomon species.
Pilsbrycharopa timorensis (B. Renscu, 1935)
Charopa brunnescens timorensis B. Renscu, 1935,
Sitz.-Ber. Ges. naturf. Freunde, Berlin, 1934: 323,
fig. 7 — West Timor; Sotem, 1958, Arch f. Mol-
lusk. 87 (1-3): 25.
Description: Shell larger than average, with slightly less
than 4 normally coiled whorls. Apex and spire slightly
and evenly elevated, body whorl descending a little more
rapidly, H/D ratio 0.541. Apical whorls 14, early sculp-
ture eroded, last one-third whorl with very low radial ribs.
In umbilicus a faint trace of spiral cording visible on first
nuclear whorl. Remaining whorls with irregular, low,
protractive radial ribs whose interstices are less than twice
their width. Microsculpture almost totally eroded, occa-
sionally traces of fine microradials and even finer micro-
spirals visible, 3 to 4 microradials between each pair of
major ribs. No trace of secondary spiral cording. Sutures
moderately impressed, whorls flattened laterally above
evenly rounded periphery and on basal margin.’ Umbili-
cus widely open, cup-shaped, contained 2.16 times in the
diameter, margins rounded. All color leached from shell.
Aperture ovate, flattened basally and laterally above peri-
phery, inclined about 30° from shell axis. Height of holo-
type 2.01 mm, diameter 3.7 mm.
Holotype: Timor: Nenas at 1000 m to 2000 m elevation.
Zoologisches Museum der Humboldt Universitat, Berlin.
Remarks: Unfortunately the holotype and only known
specimen was damaged in an attempt at cleaning heavy
incrustations. The shell is very worn and obviously collec-
ted dead. It differs from the type lot of Pilsbrycharopa
brunnescens in having a much wider umbilicus and in
lacking secondary spiral cording, although unquestionably
it is closely related. The difference in umbilical size is
large enough to warrant specific separation following the
pattern in similar situations found among Pacific Island
species.
Pilsbrycharopa brunnescens (MOLLENDoRFF, 1892)
Patula (Discus) brunnescens MOLLENDORFF, 1892,
Nachr.-Bl. dtsch. malak. Ges. 24: 87 — Tenimber
Islands; Sotem, 1958, Arch. f. Mollusk. 87 (1-3):
2a,
Diagnosis: Shell relatively large, diameter 3.82 - 4.08 mm
(mean 3.95 mm), with 44 to 44 normally coiled whorls.
Apex barely emergent or slightly elevated, lower whorls
descending slightly, body whorl more rapidly, H/D ratio
0.546-0.568 (mean 0.559). Apical whorls 13 to 14, sculp-
ture on early portion of narrow, somewhat wavy spiral
ribs, 15 or 16 in number, with faint radial ribs and a
development of strong, broadly rounded radial ribs on the
last quarter to half whorl. Postnuclear whorls with promi-
nent, lamellar, protractively sinuated radial ribs, 82-98
(mean 89.2) on the body whorl, whose interstices are 2 to
4 times their width. Ribs/mm 6.77-7.77 (mean 7.19).
Microsculpture of fine, regularly spaced radial riblets,
crossed by much finer, more crowded spiral riblets with a
secondary sculpture of moderately prominent spiral cords
whose interstices are about 3 to 4 times their width.
Sutures deep, whorls strongly rounded above, somewhat
flattened laterally above periphery and on basal margin.
Umbilicus relatively narrow, “V-shaped, regularly de-
coiling, contained 4.00-4.57 times (mean 4.31) in the dia-
meter, margins rounded. Color light reddish yellow horn
without darker flammulations. Aperture sub-circular,
slightly flattened laterally above periphery, on basal, and
on columellar margin, inclined about 25° from the shell
axis.
Pilsbrycharopa brunnescens is much smaller and has
much less prominent radial elements in the apical sculp-
ture than does P nigrofusca; P. kobelti is much smaller
and has a wider umbilicus; PR timorensis is much more
widely umbilicated..
Description: Shell large, with slightly more than 44 nor-
mally coiled whorls. Apex and spire slightly and evenly el-
evated, body whorl descending more rapidly, H/D ratio
0.556. Apical whorls 13, sculpture mainly eroded with
prominent, broadly rounded radial ribs crossed by finer,
more widely spaced spiral ribs visible on the last third. Post-
nuclear whorls with prominent, lamellar, narrow, protrac-
tively sinuated radial ribs, 86 on the body whorl, whose
interstices are 2 to 4 times their width. Microsculpture
mostly eroded, where visible consisting of regularly spaced
radial riblets crossed by much finer and more crowded
spiral riblets with a secondary sculpture of moderately
prominent, fairly widely spaced spiral cords. Sutures deep,
whorls strongly rounded above, slightly flattened laterally
above periphery and on basal margin. Umbilicus rela-
tively narrow, “V”-shaped, regularly decoiling, contained
4.00 times in the diameter. Color light reddish yellow
horn. Aperture sub-circular, inclined about 25° from the
shell axis. Height of lectotype 2.27 mm, diameter 4.08 mm.
Lectotype: Tenimber Islands. Natur-Museum Sencken-
berg, Frankfurt number 165692.
Range: Tenimber Islands and New Guinea.
Material: Tenimber Islands (4 specimens, SMF 165692-
Vol. 12; No. 3
4). Papua:- Vikaiku, Angabunga River, inland of Hall
Sound (30 specimens, MHNG, FMNH 159264).
Remarks: The type set from Timor Laut and the shells
from Vikaiku agree very well in shape, color and major
sculptural features. Both average between 15 and 16
spiral apical cords and their rib counts are essentially
identical. The slightly smaller size and lower whorl count
of the Vikaiku shells may reflect either bias for large size
in the type set remnant studied or, more probably, fewer
gerontic individuals in the former set. A possibly signifi-
cant difference is the complete absence of secondary
spiral cording in the Vikaiku shells, while it is very prom-
inent in the Tenimber sample. I consider the available
evidence too fragmentary to warrant separation, but on
the basis of variation patterns seen in other taxa I suspect
that the two are distinct.
Pilsbrycharopa brunnescens is intermediate between P
kobelti and P. nigrofusca in size and sculpture, but differs
from both in its narrower umbilicus and higher spire. The
radial apical sculpture is almost exactly intermediate in
character.
Pilsbrycharopa nigrofusca (E. A. Situ, 1896)
Charopa nigrofusca E. A. Smiru, 1896, Journ. of Ma-
lac. 5 (2): 18; plt. 2. figs. 10-12 — German New
Guinea.
Pilsbrycharopa papuana SotEeM, 1958, Arch. f. Mol-
lusk. 87 (1-3): 24-25; plt. 3, figs. 2-6 —Kon-
stantinhafen, New Guinea.
Pilsbrycharopa nigrofusca (E.A.SmrrH), SoLem,
1958, Arch. f. Mollusk. 87 (1-3): 25.
Diagnosis: Shell very large, diameter 6.14 - 6.93 mm
(mean 6.43 mm), with 44 to 44 relatively loosely coiled
whorls. Apex barely emergent, spire flat, body whorl des-
cending slightly, spire protrusion about one tenth body
whorl width, H/D ratio 0.439 - 0.489 (mean 0.462). Api-
cal whorls 1%, sculpture of fine, relatively crowded spiral
ribs, crossing lower, quite broadly rounded, somewhat re-
tractive radial ribs which are covered with very fine,
closely spaced radial riblets occasionally visible on a well-
preserved specimen. Postnuclear whorls with prominent,
“V-shaped, strongly protractively sinuated radial ribs,
114-132 (mean 124.3) on the body whorl, whose inter-
stices are 2 to 5 times their width. Microsculpture a lattice
of coequal radial and spiral riblets, equally spaced, 7 to 15
microradials between each pair of major ribs. Sutures
moderately impressed, whorls strongly rounded above,
greatly flattened laterally above periphery and slightly
below with rounded periphery and basal margin. Umbili-
cus broadly “V”-shaped, regularly decoiling, contained
THE VELIGER
Page 247
3.31 - 3.76 times (mean 3.45) in the diameter. Color
reddish yellow brown without darker maculations, apex
lighter in tone. Aperture sub-circular, strongly flattened
above periphery with slightly deflected lip and slightly
flattened lower lateral margin and umbilical edge, inclined
almost 25° from the shell axis.
The flat spire, large size, wide umbilicus, loosely coiled
whorls and microreticulated sculpture immediately sepa-
rate Pilsbrycharopa nigrofusca from the other species of
Pilsbrycharopa. Species of similar size, P. gressitti and P
schneideri, have very small umbilici and reduced radial
sculpture.
Description: (nigrofusca) Shell very large, with slightly
more than 44 relatively loosely coiled whorls. Apex and
spire flat, last whorl descending slightly, H/D ratio 0.457.
Apical whorls 13, sculpture of about 20 fine spiral ribs,
crossing much lower, broadly rounded radial ribs with a
fine secondary sculpture of radial riblets visible on lower
portion. Postnuclear whorls with low but prominent, “U”-
shaped, strongly protractively sinuated radial ribs, 114 on
the body whorl, whose interstices are 2 to 5 times their
width. Microsculpture a lattice of co-equal radial and
spiral riblets. Sutures deep, whorls sharply rounded above,
greatly flattened laterally above periphery and slightly
flattened below. Color reddish yellow brown without
darker maculations, apex lighter in color. Umbilicus broad-
ly “V-shaped, regularly decoiling, contained 3.36 times
in the diameter. Aperture subcircular, strongly flattened
laterally above periphery, slightly flattened laterally be-
low periphery and on columellar margin, inclined about
25° from the shell axis. Height of lectotype 2.81 mm,
diameter 6.15 mm.
(papuana) Shell very large, with 43 relatively loosely
coiled whorls. Apex and spire flat, body whorl moderately
descending, H/D ratio 0.443. Apical whorls 14, sculpture
of fine spiral ribs, about 23 in number, reticulated by
slightly finer radial ribs on the early portion with an
intrusion of broadly rounded radial ribs on the last one-
third whorl. Postnuclear whorls with rounded, prominent,
protractively sinuated radial ribs, 116 on the body whorl,
whose interstices are 2 to 3 times their width. Microsculp-
ture a lattice of co-equal radial and spiral riblets in a
somewhat waved pattern. Sutures deeply impressed,
whorls evenly rounded at shoulder, strongly flattened lat-
erally above periphery and on basal margin, with evenly
rounded periphery. Color dark reddish yellow horn, apex
somewhat lighter in tone. Umbilicus widely opened, “V”-
shaped, regularly decoiling, contained 3.42 times in the
diameter. Aperture large, ovate, flattened laterally above
periphery and slightly on basal margin, inclined about 25°
from the shell axis. Height of holotype 3.08 mm, diameter
6.93 mm.
Page 248
Holotype of papuana: New Guinea: Constantinhaven.
Natur-Museum Senckenberg, Frankfurt number 158151/1
ex-Mollendorff.
Lectotype of nigrofusca: New Guinea: Konstantinhaven.
British Museum (Natural History) number 96.6.1.21.
Range: Known only from Konstantinhaven, New Guinea.
Paratypes: B. M. (N.H.) 96.6.1.22 (nigrofusca) ; SMF
158182/3, FMNH 63527, UMMZ 141799 (papuana).
Remarks: Inspection of the types of Charopa nigrofusca
SmiTH, 1896, showed that Pilsbrycharopa papuana SOLEM,
1958, is a synonym, probably based upon material from
the type lot of the former. The original figures of C. nigro-
fusca are incorrect in proportions and led to the erroneous
assumption that the two taxa were separable.
The large size and flattened form of Pilsbrycharopa
nigrofusca are the obvious features separating it from P
brunnescens. This enlargement was not achieved by ad-
ding whorls, but simply through overall increase. Rib
spacing is slightly wider, but only to the extent propor-
tional to size change. In having a slightly lower spire and
proportionately wider umbilicus, P nigrofusca shows a
characteristic linked change. In this instance, it is caused
by depressing the plane of coiling and flattening of the
whorls. Pilsbrycharopa brunnescens from Vikaiku has the
spire protrusion/body whorl width index average 0.098,
while in P nigrofusca it is 0.105 (see Table 2). The spire
of P. brunnescens appears much higher, but the measured
protrusion is only slightly less. Body whorl width averages
1.18 mm in P. brunnescens and is 1.88 mm in P. nigrofusca.
There is thus an increase of 77% in mean diameter be-
tween the species (3.63 in Vikaiku PR brunnescens to 6.43
in RP nigrofusca) but only a 59% increase in body whorl
width. This is caused by more marked compression of the
whorls in P. nigrofusca.
Much more significance can be attached to the greater
radial element in the apical sculpture of Pilsbrycharopa
nigro fusca.
Pilsbrycharopa densecostulata (THIELE, 1928)
Charopa densecostulata TutELE, 1928, Zool. Jahrb.,
Syst. 55: 127; plt. 5, fig. 14 -—29km unterhalb
Maeanderberg (Upper Sepik River), New Guinea.
Pilsbrycharopa densecostulata (TuHrte), SoLe,
1958, Arch. f. Mollusk. 87 (1-3): 25.
Remarks: I have not seen the single known example of
this species, but the original description and figures leave
no doubt as to its affinities. The apical sculpture, shape,
umbilical form and radial sculpture all appear to be inter-
THE VELIGER
Vol. 12; No. 3
mediate between the features of species such as Pilsbry-
charopa brunnescens and the very specialized PR renschi
or P. schneideri.
Maeanderberg is a classic collecting area on the Upper
Sepik River, Sepik District, New Guinea located past the
great bend of the Sepik quite near the border of West
Irian at about 4°10’ S, 141°25’ E.
Pilsbrycharopa gressitti SOLEM, spec. nov.
(Figures 1d to 1f; 2f to 21)
Diagnosis: Shell very large, diameter 6.27 mm - 7.32 mm
(mean 6.76 mm), with 3 to 44 normally coiled whorls.
Apex and spire slightly to moderately and evenly elevated,
spire protrusion averaging one-ninth body whorl width,
last whorl descending more rapidly, H/D ratio 0.559-0.642
(mean 0.585). Apical whorls slightly more than 14,
sculpture of about 15 - 18 fine and widely spaced spiral
cords crossing much higher and more widely spaced radial
ribs that become more crowded near end of apex. Post-
nuclear whorls with narrow, low rounded radial ribs, in
fresh material with fine periostracal extensions, that be-
come extremely crowded and indistinct by middle of body
whorl. Microsculpture occasionally distinguishable as very
fine and crowded radial riblets, much finer and more
crowded spiral riblets, with a secondary sculpture of low,
very broad spiral cords whose interstices are usually less
than twice their width. Sutures moderately impressed,
whorls strongly rounded above, compressed laterally above
periphery and on basal margin, with evenly rounded peri-
phery and columellar margins. Umbilicus very narrow,
“V-shaped, slightly decoiling, partly covered in adults by
rolled reflection of columellar lip, contained 9.27 - 12.47
times (mean 10.74) in the diameter. Aperture ovate,
compressed laterally above periphery and on basal margin,
inclined about 25° from shell axis.
Pilsbrycharopa gressitti has only a slightly elevated
spire, a narrow umbilicus, and is quite large. Species of
similar size differ in having a minute or closed umbilicus
and grossly elevated spire (P. schneideri) or a wide umbili-
cus and regular ribbing (P nigrofusca). None of the other
Pilsbrycharopa exceed 4mm in mean diameter.
Description: Shell very large, with 3$ normally coiled
whorls. Apex and spire very slightly and evenly elevated,
last whorl descending much more rapidly, H/D ratio
0.563. Apical whorls more than 14, early portion wom,
lower part with 16 narrow spiral cords, whose interstices
are 5 to 6 times their width, with much higher, narrow,
rounded, widely spaced radial ribs, whose interstices at first
are 4 to 5 times their width, becoming more crowded near
Vol. 12; No. 3 THE VELIGER Page 249
ES
AALLLEL Biri
abc
-——
Figure 1
a-c: Holotype of Paryphantopsis dauloensis SoLEM, spec. nov. d-f: Holotype of Pilsbrycharopa gressitti SOLEM, spec. nov.
Daulo Pass, Eastern Highlands, New Guinea, Bernice P Bishop Daulo Pass, Eastern Highlands, New Guinea, Bernice P. Bishop
Museum Museum.
Scale lines equal 1 mm
Page 250 THE VELIGER ; Vol. 12; No. 3
Page a ——————————————a
Figure 2
a-e: Paryphantopsis dauloensis SoLEM, spec. nov. Type lot. f-i: Pilsbrycharopa gressitti SoLEM, spec. nov. Type lot.
Daulo Pass, Eastern Highlands, New Guinea. a - pallial region; Daulo Pass, Eastern Highlands, New Guinea. f - pallial region;
b-genitalia; | c-d-detail of epiphallic diverticulum in two spe- g-h- terminal genitalia; i- interior of. penis
cimens;_ e - interior of penis Scale lines equal 1 mm
Dissections deposited in Bernice P. Bishop Museum
Vol. 12; No. 3
end of apex. Postnuclear sculpture of low, rounded, radial
ribs with slight periostracal extensions, becoming very
crowded lower on spire, indistinguishable on body whorl.
Microsculpture as in diagnosis. Sutures impressed, whorls
strongly rounded above, compressed laterally above peri-
phery and on basal margin, with evenly rounded peri-
phery. Umbilicus very narrow, “V-shaped, slightly de-
coiling, contained 12.44 times in the diameter, partly
narrowed at aperture by columellar lip reflection. Aper-
ture ovate, compressed laterally above periphery and
slightly on basal margin, inclined about 30° from shell
axis. Height of holotype 4.12 mm, diameter 7.32 mm.
Holotype: New Guinea: Daulo Pass (ca. 6°03’S, 145°10’
W), Eastern Highlands, at about 8200 feet elevation.
Collected on June 14, 1955 by J. Linsley Gressitt. Bernice
P. Bishop Museum.
Material: Daulo Pass (7 specimens, BPBM).
Remarks: The very characteristic apical sculpture is per-
fectly captured in Figure 1 d. In contrast to that of Pils-
brycharopa nigrofusca (see SoLeM, 1958, plt. 3, fig. 2) the
radial elements of the apical sculpture are narrow and
distinctly higher than the spiral cords. Pilsbrycharopa
schneider: has slightly greater accentuation of the radial
elements.
Great pleasure is taken in naming this species after
its collector, J. Linsley Gressitt, whose work on the Pacific
Island fauna has stimulated such interest in the Pacific
biota.
Description of soft parts: Foot broad, truncated anterior-
ly, tail slightly tapering, bluntly rounded behind. Sole
transversely corrugated in preservative, without longitu-
dinal zonation. Pedal grooves deeply impressed, high on
side of foot, suprapedal smaller than pedal, uniting above
tail, no caudal horn or middorsal groove present. Slime
network irregularly rectangular, units much smaller than
lateral divisions below pedal grooves. Head and ommato-
phores without marked peculiarities. Gonopore position
not observed because of extreme retraction in available
material. Body color light on sole and back of tail, an
iridescent grey on sides of foot and neck, sole yellow-white.
Mantle collar (MC) wide and thick, no glandular ex-
tension onto mantle roof. Anus (A) opening just inside
pneumostome, distinctly anterior of external ureteric pore
(KX). Pallial region (Figure 2 f) measures 6.25 mm from
edge of mantle collar to posterior end of kidney, distance
from anterior edge of kidney to mantle collar about 2.6
mm. Lung roof clear of granulation. Kidney (K) bilobed,
about 3.4mm long, slanting downward from _parietal-
palatal margin after anterior third of rectal lobe, tapering
posteriorly under intestinal loops. Ureter (KD) typical,
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Page 251
narrow strip of lung roof visible between arms of ureter
which are narrow at point of reflection. Heart (H) large,
two thirds length of kidney. Principal pulmonary vein
(HV) very inconspicuous, without major branching vis-
ible. Hindgut (HG) departing from kidney about 3.65
mm behind edge of mantle collar. Apical genitalia not
seen. Prostate (DG) of numerous slender acini opening
into groove on inner surface of upper uterine chamber.
Uterus (UT) bipartite, upper chamber very slender and
thin walled, lower chamber swollen, biscuit-shaped, with
thick glandular walls. Vas deferens (Figure 2, g-h) very
large and glandular at first, narrowing to a slender tube
before penioviducal angle, reflexing up to enter bulbous
epiphallic head. Epiphallus (E) with swollen head nar-
rowing after an anterior diverticulum (EL) followed by
a slender tube with longitudinal pilasters leading to penis
head. Penial retractor (PR) arising on diaphragm, rather
long, inserting longitudinally on coiled shaft of epiphallus
below epiphallic diverticulum. Penis (P) about 2mm
long, elongately ovate, tapering from bulbous head down
to slender atrium. Internally (Figure 21) with short lon-
gitudinal pilasters clustering around epiphallic pore, main
portion with modified stimulatory pad and accessory pil-
asters, walls very dense and muscular. Atrium (Y) very
short and slender. Free oviduct (UV) short, a narrow
tube opening into very short vagina lateral to spermathe-
cal insertion, without clearly defined internal structures
other than longitudinal grooves. Spermatheca (S) with
ovoid head lying just above prostate-uterus, slender shaft
lightly bound to surface of prostate, approximately doub-
ling in size just before inserting on penioviducal angle.
Vagina (V) a short thick-walled, wide tube with heavy
muscular walls. Free muscle system not studied, except to
note typical passage of right ommatophoral retractor
through penioviducal angle. Jaw fragmented in mounting,
separate plates, about 3 times as long as wide on outer
portions, central area fused. Radula with central about lu
narrower than 1* lateral, mesocone extending slightly be-
yond tip of basal plate. Laterals 12-13, 1* with basal
plate about 13m wide and 16 long, mesocone extending
up 6u beyond edge of basal plate. Ectocone and endocone
equal in size on 1“ lateral, endocone increasing more
rapidly than ectocone on outer laterals, basal plate be-
coming shorter and wider. Shift to marginals in one or two
teeth by sharp reduction in mesoconal length, less dram-
atic size increase in ectocone and endocone, shift in basal
plate from almost square to shorter than wide. Marginals
9 to 10, outer with basal plate width twice length, cusps
becoming shorter, ectocone split by 8" marginal
marginals often only with mesoconal remnant. (Based on
3 fragmentary specimens from the type set. They had
been extracted previously with the apical parts remaining
in the shells.)
Page 252
Pilsbrycharopa renschi (Franc, 1952)
Charopa vicina B.RENscH, 1930 (not PRESTON
1907), Zool. Anz. 89 (3-4): 87 — Batoe Doelang,
Sumbawa and Rana Mésé, Flores, Lesser Sunda
Islands; B. RenscuH, 1932, Zool. Jahrb., Syst. 63:
102; plt. 3, fig. 34; Sorem, 1958, Arch. f Mollusk.,
87 (1-3): 25.
Charopa renschi Franc, 1952, Bull. Soc. France 77
(1): 78 — New name for Charopa vicina RenscuH,
1930 not Preston, 1907.
Description: Shell larger than average, with slightly more
than 54 tightly coiled whorls. Apex and spire markedly
and evenly elevated, spire protrusion about two fifths
body whorl width, last whorl descending very slightly,
H/D ratio 0.750. Entire shell encrusted and sculpture
partly destroyed by fungal attack. Apex with both radial
and spiral sculpture, details not detectable. Postnuclear
whorls with fine, very crowded, vertical radial ribs, about
230 on the body whorl, that become retractive near umbil-
ical closure. Microsculpture a lattice of radial and spiral el-
ements, the latter weaker. All sculpture fainter below peri-
phery of body whorl. Sutures not very deep, whorls
strongly and evenly rounded above, becoming flatly
rounded on outer margin. Umbilicus completely closed
by reflection of lip. Color yellow horn with faint reddish
tinge. Aperture crescentic, slightly more strongly rounded
above and on umbilical margin. Columellar lip thickened
and reflected. Height of holotype 2.86mm, diameter
3.82 mm.
Holotype: Sumbawa, Batoe Doelang, west part of island,
at 1000 to 2000 m elevation. Zoologisches Museum der
Humboldt Universitat, Berlin.
Paratype: Flores, Rana Mésé at 1 200 - 1 300 m elevation
(SMF 5733).
Additional Material: Sumbawa, Tamboka at 3000 feet
elevation (BMNH 98.10.25.184).
Remarks: The juvenile paratype is badly broken, has the
apical whorls missing, and a narrowly open umbilicus.
Although the juvenile (32 whorls) from Tamboka was
somewhat worn, it showed that the apical sculpture con-
sists of fine, very crowded spiral riblets with smaller,
slightly lower radial riblets and an intrusion of low
rounded radial ribs on the lower part of the apex. This
serves to relate P renschi to Pilsbrycharopa despite the
quite different shape. Differences from P baliana are given
under that species.
THE VELIGER
Vol. 12; No. 3
Pilsbrycharopa baliana (B. RENscu, 1930)
Charopa baliana B. ReNscH, 1930, Zool. Anz., 89
(3-4): 86; fig. 14 -— Gitgit and Batoeriti, Bali,
Lesser Sunda Islands; B. ReENscu, 1932, Zool.
Jahrb., Syst. 63: 102; Sorem, 1958, Arch. f. Mol-
lusk., 87 (1-3): 25.
Description: Shell larger than average, with 5} very
tightly coiled whorls. Apex and spire markedly and evenly
elevated, body whorl not descending more rapidly, spire
protrusion a little less than two fifths body whorl width,
H/D ratio 0.729. Apical and early postnuclear whorls
with sculpture eroded. Lower whorls with thin, vertical,
radial ribs, 151 on the body whorl, whose interstices are
3 to 5 times their width. Ribs become retractively sinuated
upon basal umbilical area. Ribs/mm 12.39. Microsculp-
ture a lattice of radial and spiral riblets, the latter dis-
tinctly finer. Both macro- and microsculpture weaker be-
low periphery of body whorl. Sutures deep, whorls strongly
rounded above slightly angled periphery, compressed lat-
erally, flatly rounded to basal margin, which is strongly
rounded. Umbilicus a narrow crack, contained about 20
times in the diameter. Color reddish-yellow horn without
flammulations. Aperture crescentic, strongly rounded on
umbilical and upper palatal margins, parallel to shell
axis. Columellar margin thickened. Height of holotype
2.83 mm, diameter 3.88 mm.
Holotype: Bali, Gitgit at 500- 600m elevation. Zoolo-
gisches Museum der Humboldt Universitat, Berlin.
Paratype: Bali, Batoeriti at 800 m elevation (SMF 5734).
Remarks: The paratype differed only in being slightly
less elevated (H/D ratio 0.698) and a trifle smaller (dia-
meter 3.82 mm). The whorl count was identical. Unfor-
tunately, the apical sculpture was eroded.
The very similar Pilsbrycharopa renschi (FRANG, 1952)
differs in having a closed umbilicus when adult, finer and
more crowded radial sculpture (ca. 230 ribs on the body
whorl), a slightly less obtuse spire angle, and shallower
sutures.
Pilsbrycharopa schneideri (1. RENscu, 1937)
Charopa schneideri 1. RENscH, 1937, Arch. f. Natur-
gesch., N. FE, 6 (4): 587 - 588; figs. 29 - 31 (shell,
genitalia, radula) — Malkong-Bach, Karlei, Ma-
tong, Lomal, Patagun, Ulamona and Insel Lolo-
bau, New Britain, Bismarck Archipelago.
Vol. 12; No. 3
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Page 253
Table 3
Variation in Groups “B-E” Species of Pilsbrycharopa
° °
6§3 | 4 2g
eh E “ 3 e E Z E
EB 8 3 & a 3 E ~
pane EY ae a Fe 5 Q
P. densecostulata 0 = 23 3.75 0.613 32 0.67 5.60
(THIELE)
P. gressitti SoLeM, GT RS 3.95+0.136 | 6.76+0.147 | 0.585+0.0131 44 0.60+0.030 | 10.74+0.518
spec. nov., Daulo Pass (3.53-4.44) | (6.27-7.32) | (0.559-0.642) | (33-44) (0.52-0.72) | (9.27-12.47)
P. baliana (B. Renscw) | ALPS 2.75 3.85 0.714 5} 0.16 24.4
Bali
P. renschi (FRANC) 1 230 19.17 3.82 0.750 54 closed -
Sumbawa
P. schneideri (I. ReNscH) 22 = 3.99+0.148 | 5.95+0.151 | 0.669+0.0113 4h crack to over
Malkong (3.01-5.82) | (5.10-7.91) | (0.588-0.774) | (43-6) closed 20
Vikaiku 1 | - | 4.38 0.705 44 crack -
Buja Kori a re s 3.36+0.084 | 5.52+0.108 | 0.608+0.0161 434 crack =
(3.20-3.60) | (5.23-5.69) | (0.563-0.632) | (44-44)
Diagnosis: Shell very large, diameter 5.10 to 7.91mm
(mean 5.95mm), with 43 to 6 rather tightly coiled
whorls. Apex and spire moderately to very strongly ele-
vated, spire not rounded above, spire protrusion more
than one third body whorl width, H/D ratio 0.588 - 0.774
(mean 0.669). Apical whorls 13 to 13, sculpture of curved,
slightly retractive, close-set radial ribs, whose interstices
are about equal to their width, plus distinctly smaller
spiral ribs. In worn specimens this approximates a “pit-
ted” sculpture. Postnuclear whorls with low, rather irreg-
ular, strongly protractively sinuated radial ribs, about 145
on the body whorl, whose interstices are 3 to 6 times their
width. Microsculpture of rather coarse radial riblets with
exceedingly fine spiral reticulation and a secondary sculp-
ture of broad, widely spaced spiral cords. Where the
spiral cords cross the radial ribs and riblets, a “beaded”
effect results. Suture impressed, whorls slightly shouldered
above, outer and basal margins evenly rounded. Umbilicus
completely closed, a slight crack, or a very narrow opening
contained more than 20 times in the diameter. Color light
yellow-white with frequent radial reddish flammulations
that coalesce over much of the shell spire. Aperture sub-
circular, slightly shouldered above, with evenly rounded
outer margins, basal margin extended flatly to columella.
Columellar lip thickened by white callus, reflected and
twisted to close umbilicus, adults with a distinct columel-
lar-basal angle. Aperture inclined about 15° from shell
axis.
Table 4
Size and Shape Variation in Paryphantopsis
Specimens
Examined
N umber of
P. platycephala 3.04 + 0.042
(2.94-3.14)
3.41 +0.074
(3.01-3.79)
P. fultoni
Diameter
4.93+0.188 | 0.618+0.0171 +
(4.71-5.49) | (0.571-0.653) | (23-22)
8.52+0.127 | 0.401+0.0098 28
(7.97-9.28) | (0.354-0.434) (28-3)
Page 254
THE VELIGER
Vol. 12; No. 3
Pilsbrycharopa schneideri is very large, with a nearly
closed or closed umbilicus, protruding spire, and altered
radial sculpture. Pilsbrycharopa gressitti and P. nigrofusca
have much finer sculpture, open umbilici and much less
elevated spires. Other Pilsbrycharopa are much, much
smaller in size.
Paratypes: New Britain: (ZMB, FMNH
146031).
Malkong
Material: New Britain: Malkong (30 specimens, ZMB,
FMNH 146031). Vikaiku, village on Angabunga or St.
Joseph River, inland from Hall Sound (2 specimens, MH
NG collected by Lamberto Loria in July 1892) ; Bujakori,
village along Kemp Welch River, northeast of Rigo (9
specimens, MHNG, FMNH 159264, collected by Lam-
berto Loria in August 1890).
Remarks: Pilsbrycharopa schneideri differs from the other
New Guinea species in its very elevated spire, having the
radial apical sculpture primary, the flared baso-columellar
lip, closed or cracked umbilicus, irregular radial ribbing,
and very strong secondary spiral cording. The Indonesian
P. renschi and P. baliana are very similar in general ap-
pearance, but have typical endodontid radial ribs and a
less specialized apical sculpture.
Paratypes from Malkong came from river drift piles
and showed considerable variation in size and shape (Table
3). Both the Vikaiku adult and 4 Bujakori examples fall
within the range of variation shown by this set. I can see
no sculptural differences between the shells and do not
hesitate to consider them conspecific. Adult shells can be
recognized by the angled baso-columellar margin and
generally closed umbilicus. Juveniles have a rounded mar-
gin and slightly open umbilicus.
I. Renscu (loc. cit.) gave outline figures of the termi-
nal genitalia and radular teeth. Apparently there is no
epiphallic diverticulum as in Pilsbrycharopa gressitti, and
the illustrated vas-deferens-epiphallic union seems pecul-
iar. Without study of the penis interior and verification
of the epiphallic diverticulum absence, I would hesitate to
separate this species from Pilsbrycharopa.
Paryphantopsis THIELE, 1928
Zool. Jahrb., Syst. 55: 125 - 126.
Medium sized to very large Charopinae in which the
apical sculpture varies from a lattice of co-equal radial
and spiral riblets to distinctly pitted surface. Whorls very
loosely coiled, generally about 3, rarely more. Spire nor-
mally slightly raised, flat in many species. Umbilicus nor-
mally closed, narrowly open laterally only in Paryphantop-
sis dauloensis and P. sculpturata. Radial sculpture reduced
to periostracal fringes in most taxa, remnants of major
radials in P striata and P. louisiadarum. Whorls keeled in
P elegans and P. fultoni, rounded in other species. Anato-
my known only in one species. Pallial cavity shortened,
hindgut and kidney separating early, arms of ureter widely
spread, pulmonary vein markedly branched. Terminal
genitalia with short penial retractor muscle, tapering short
penis with vergic papilla and large corrugated pilasters,
a long epiphallus and epiphallic diverticulum.
Type species: Flammulina (Paryphantopsis) lamelligera
THIELE, 1928 by OD.
Except for the material of Paryphantopsis fultoni and
P. platycephala reported on below, no species of this
genus is known from more than 3 specimens. Variation in
4 adults of P platycephala and 9 P fultoni is summarized
in Table 4.
Obviously we have only the most rudimentary know-
ledge of intra-populational variation. This makes it impos-
sible to attempt a formal revision on the species level. I
have examined the types of all but Paryphantopsis similis
THIELE, 1928, P lamelligera Tueve, 1928 and P. globosa
(HEDLEY, 1890). Type specimen descriptions are pre-
sented for species described prior to the work of van
BENTHEM JuTTING (1964, pp. 13-17), but her species
are well defined and easily identifiable.
Table 5 summarizes available data on species characters.
The taxa show differing combinations of relatively few
changes in character states. Paryphantopsis fultoni and P
elegans are immediately recognizable because of their
sharp peripheral keels; P lowisiadarum and P. striata by
the lack of periostracal fringing on the ribs. Otherwise
the differences are in size, spire and apex elevation, body
whorl descension and presence or absence of an umbilical
chink. These are comparatively minor changes and the
general appearance of Paryphantopsis species is very
similar.
No specimens have been taken in the fairly well col-
lected Vogelkop or Biak Island areas of Irian. All known
species have been collected between about 139° E near the
Idenburg River of Irian to Rossel Island, Louisiades,
Papua. Specimens of Paryphantopsis fultont, P. platyceph-
ala and a shell tentatively identified as P globosa have been
taken near Moroka on the Laloki River, Papua; P. filosa
and P. sculpturata above the Idenburg River; P Jatior and
P. arcuata in the Star Mountains of Irian. Apparently
there is some sympatry or near sympatry within the genus.
Members of the last two species pairs were taken at dif-
ferent elevations, but within a few miles of each other.
Judgment of distribution and speciation patterns must
Vol. 12; No. 3 THE VELIGER Page 255
Table 5
Type or Mean Adult Characters in Paryphantopsis
S 3 $8
se] § A 3
mT: z Be Si | s
= a oa $3 4
Se Sel | Ss oe ae 5
Paryphantopsis (P)
filosa 2.7 0.750 Ton Raised Strongly Closed
eed
Pyamaca spon] Clas
arcuata 8 | 0646 | 3 Raised || Closed
sculpturata 3.3 0.705 Slightly Open
Raised
similis 0.700 Raised Closed
platycephala
Type 2.7 Flat Slight Closed
Moroka 3.04 Flat Slight Closed
dauloensis 3.26 Raised Yes uk Open
eile
elegans 2.55 Raised No Closed
latior Flat | Rapid | Closed
lamelligera Raised Yes Closed
striata 5.40 Raised Rapid Closed
fultoni 3.40 Slightly No Closed
Raised
loutsiadarum 6.50 Flat Yes Closed
P. (Gallodema)
globosa 4.44 | 5.95 Raised Raised Yes Crack
(Moroka Juvenile)
Original Description | 10.0 | 12.0 Raised Raised Yes Open
await much larger and geographically more diverse
samples.
Recognition of species group is equally premature, with
only the single species, Paryphantopsis dauloensis, dis-
sected. An exception concerns the position of the shell
described as Rhytida globosa Hepiey, 1899. IREDALE
(1941, p. 92) proposed a generic name, Gallodema, and
suggested it might be “an aberrant Zonitid” or a pary-
phantid. He also described the genus Illonesta for Pary-
phanta louisiadarum MOLLENDORFF, 1899, having over-
looked the description of Paryphantopsis THIELE, 1928.
As can be seen from Table 5, HEpLEy’s species differs
greatly in size, whorl count and major sculpture from the
typical Paryphantopsis. In apical sculpture, however, it is
intermediate between Paryphantopsis and the more speci-
alized Pilsbrycharopa, and the largest Paryphantopsis are
nearer in size than the largest Pilsbrycharopa. Pending
availability of further material for dissection, I prefer to
classify HEpLEy’s species as a Paryphantopsis, but use
Gallodema as a subgeneric name.
No key to the species is attempted, since simple com-
parison of the measurements and variable shell features
with the data in Table 5 will indicate which are the most
similar morphotypes.
Species are listed in ascending order of size. Comments
are restricted to addition of new or supplemental data.
For most species I have included only references to pre-
vious literature.
Paryphantopsis (Paryphantopsis) filosa vAN BENT-
HEM JuTTING, 1964
Paryphantopsis filosa VAN BENTHEM JUTTING,
1964, Nova Guinea, Zool., 26: 14-15; figs.
Page 256
THE VELIGER
Vol. 12; No. 3
16-19 — Araucariakamp at 800 m elevation,
south of Idenburg River, near Vlakke Peak,
north side Snow Mts., Irian (ca. 139°10’ E,
3°30’S).
Paryphantopsis (Paryphantopsis) pygmaea (Bavay,
1908)
Helicarion pygmaeus Bavay, 1908, Nova Gui-
nea, Zool., 5: 286 - 287; plt. 14, figs. 14, a, b, c
— Cyclopis montern Novae Guineae (= Cyclops
Mts., near Hollandia, West Irian, ca. 140°35’
1%, DBO So
Paryphantopsis pygmaea (BAvAy), VAN BENT-
HEM JutTTING, 1964, Nova Guinea, Zool., 26:
13 - 14.
Paryphantopsis (Paryphantopsis) arcuata VAN BENT-
HEM JuTTING, 1964.
Paryphantopsis arcuata VAN BENTHEM JUT-
TING, 1964, Nova Guinea, Zool., 26: 16; figs.
24-27 — Nimdol, bivak 36 at 1220 m eleva-
tion, Star Mountains, Irian (ca. 140°48’ E,
4°54’S).
Paryphantopsis (Paryphantopsis) sculpturata vAN
BENTHEM JuTTING, 1964
Paryphantopsis sculpturata VAN BENTHEM
Jurtinc, 1964, Nova Guinea, Zool., 26: 14;
figs. 12-15 -— Rotankamp and Tussenkamp
at 1100 - 1 200 m elevation, south of Idenburg
River, near Vlakke Peak, north side Snow Mts.,
Irian (ca. 139°10’ E, 3°30’ S).
Paryphantopsis (Paryphantopsis) similis (THIELE,
1928)
Flammulina (Paryphantopsis) similis THiE.e,
1928, Zool. Jahrb., Syst., 55: 127; plt. 5, figs.
12, a — Maeanderberg, Upper Sepik River,
Sepik District, New Guinea (ca. 147°40’ E,
6°30’S).
Paryphantopsis (Paryphantopsis) platycephala vAN
BENTHEM JUTTING, 1964
Paryphantopsis platycephala vAN BENTHEM
Juttine, 1964, Nova Guinea, Zool., 26: 16-17;
figs. 28-31 — Doormanpad bivak at 1410m
elevation, above Doorman River, Upper Mam-
beramo drainage, West Irian (ca. 138°30’ E,
3) es fS)))e
Material: Papua, Moroka, headwaters of Laloki River,
east of Port Moresby, Central District (10 specimens, MH
NG, FMNH 159262).
Remarks: Four adults and six juveniles from near Moro-
ka, collected by Lamberto Loria in 1893, are tentatively
referred to Paryphantopsis platycephala. They are nearest
to this species, although showing some differences from
the type in height and H/D ratio. Variation in the few
adults is summarized in Table 4.
Paryphantopsis (Paryphantopsis) dauloensis, SoLEM
spec. nov.
(Figures 1 a to 1c; 2a to 2e)
Diagnosis: Shell slightly smaller than average, diameter
5.36mm to 5.59mm (mean 5.48mm), with 3 to 34
loosely coiled whorls. Apex and spire barely to moderately
and evenly elevated, body whorl descending moderately to
much more rapidly, H/D ratio 0.577-0.612 (mean 0.595).
Apical whorls 14, rounded, sculpture of rather vague radi-
al and spiral riblets producing a weakly pitted appearance
by partial fusion and wear. Postnuclear whorls with irreg-
ular growth wrinkles, many with prolonged periostracal
extensions, frequency and prominence of extensions re-
duced on shell base. Microsculpture mostly absent, occa-
sionally visible under 96 magnification as fine radial
and slightly finer spiral riblets. Sutures deep, whorls
strongly rounded above, compressed laterally above and
below rounded periphery. Umbilicus narrowly and later-
ally open, rolled reflection of columellar lip partly con-
stricting opening. Aperture very large, subcircular, com-
pressed laterally above and below rounded periphery,
inclined about 40° from shell axis.
The open umbilicus, absence of spiral pitting on the -
postnuclear whorls, reduced fringing and weak apical
sculpture separate Paryphantopsis dauloensis from pre-
viously described species. Paryphantopsis latior VAN BENT-
HEM JUTTING has a flat apex and spire, closed umbilicus
and is almost 1 mm larger in mean diameter; P similis
has a closed umbilicus, is smaller, much higher and has
the periostracal fringes much more prominent; P sculp-
turata has an open umbilicus, prominent spiral pitting on
the lower whorls, and is almost 1 mm smaller in mean
diameter.
Description: Shell smaller than average, with 3 loosely
coiled whorls. Apex and spire slightly protruding, body
whorl descending more rapidly, H/D ratio 0.577. Apical
whorls 14, evenly rounded, surface cracked and partially
obscured by fungus, sculpture of low radial and spiral
riblets of equal size, tending towards a pitted appearance.
Postnuclear whorls with irregular radial growth wrinkles,
many with prominent periostracal extensions. Microsculp-
ture occasionally visible as fine radial and slightly finer
spiral riblets, surface usually smooth. Sutures deep, whorls
strongly rounded above, compressed laterally above and
below rounded periphery. Color reddish-brown, becoming
Vol. 12; No. 3
yellow-brown near apex. Umbilicus narrowly and later-
ally open, periostracal reflection of columellar lip covering
opening in direct bottom view. Aperture very large, in-
clined about 40° from shell axis, margin only weakly
sinuated. Height of holotype 3.09 mm, diameter 5.36 mm.
Holotype: New Guinea: Daulo Pass, Eastern Highlands
at about 8200 feet elevation. Collected by J. Linsley
Gressitt on June 13, 1955. Bernice P. Bishop Museum.
Range: Known only from the type collection.
Material: Daulo Pass (2 specimens, BPBM).
Remarks: Of the previously described Paryphantopsis,
only P sculpturata vaN BENTHEM JUTTING has a partly
open umbilicus. It is smaller, higher and has prominent
spiral sculpture on the lower whorls.
Description of the soft parts: Foot and tail shorter than
in Pilsbrycharopa. Sole broad and transversely corrugated.
Pedal grooves high on foot, deeply impressed, suprapedal
less conspicuous, both united over tail, no caudal horn or
middorsal groove present. Slime network typical. Gono-
pore position not observed. Body color iridescent yellow-
white with a very faint reddish tint (caused by preserva-
tive?). Mantle collar thick and with low lobes, but no
glandular extension onto pallial roof. Pneumostome
masked by swellings on mantle edge. Anus opening just
inside pneumostome, external ureteric pore lying right
beside anus. Pallial region (Figure 2 a) measures 2.96 mm
from edge of mantle collar to peak of kidney, distance
from anterior end of kidney to mantle collar about
1.50 mm. Lung roof clear of granulations. Kidney bilobed,
lobes almost equal in length, high in center, slanting back-
wards under intestinal looping beyond end of pallial ca-
vity. Posterior portion of kidney sharply angled down from
hindgut and parietal-palatal margin. Ureter reflexed,
complete, lung roof visible between arms of ureter, tube not
tapering, compressed at point of reflection. Heart large,
more than two thirds length of pericardial kidney arm,
slightly hidden posteriorly by margin of kidney. Principal
pulmonary vein very inconspicuous, without major branch-
ing after initial bifurcation. Hindgut departing from ure-
ter 1.4mm behind anus. Apical genitalia not seen. Pros-
tate and uterus as in Pilsbrycharopa. Vas deferens
(Figure 2b, VD) large at first, becoming very slender,
entering epiphallus at swollen head after being coiled once
around penis at muscle sheath. Epiphallus (E) with swol-
len head, long lateral diverticulum (EL), and long, coiled
tube leading to penis head. Diverticulum equal in length
to epiphallic head (Figures 2 c, 2d). Penial retractor (PR)
arising from diaphragm, very short, inserting on epiphallic
tube near entrance to penis. Penis (P) about 1.9mm
THE VELIGER
Page 257
long, club-shaped, gradually tapering, basal third a slender
tube with muscular sheath. Internally (Figure 2 e) with
short vergic papilla (PV) with central groove, sheath of
verge attached to one wall of penis, other walls with cor-
rugated longitudinal pilasters (PP) tapering into atrium.
Atrium (Y) very short, rather broad. Free oviduct (UV)
short, internally with longitudinal pilasters, opening into
vagina. Spermatheca (S) with basal portion a muscular
tube exceeding in diameter free oviduct, narrowing at base
of prostate-uterus to slender tube passing up prostate sur-
face, head club-shaped, gradually expanding, lying slight-
ly above apex of prostate-uterus. Vagina (V) greatly ex-
panded, with very thick glandular walls. Free muscle
system without unusual features. Jaw of narrow plates,
length 5 or 6 times width, fused centrally and tightly
joined out to margins. Width about 0.10 mm to 0.11 mm,
length not determined since jaw broken in mounting.
Central slightly smaller than 1* lateral, mesocone cusp
barely projecting beyond basal plate. Laterals 10 to 12,
1* about 10 wide, 13 w long, cusp projecting beyond end
of basal plate. Ectocones and endocones all smaller than
in Pilsbrycharopa. Marginals with square basal plates, 7 to
8 in number, single ectocone and endocone two thirds
length of mesocone.
(Based on two broken specimens extracted from shell and
broken off in process).
Paryphantopsis (Paryphantopsis) elegans (FuttoN, 1902)
Paryphanta elegans Futon, 1902, Ann. Mag. Nat.
Hist., (7). 9: 182 - 183 — Arva (=Aroa) River,
New Guinea.
Illonesta elegans (FULTON), IREDALE, 1941, Austral.
Zool., 10 (1): 93.
Paryphantopsis (Paryphantopsis) elegans (FuLToN),
Sotem, 1958, Arch. f. Mollusk., 87 (1-3): 23;
1959, loc. cit., 88 (4-6): 156.
Description: Shell of average size, with 3 very loosely
coiled whorls. Apex and spire slightly and evenly elevated,
last whorl not descending more rapidly, H/D ratio 0.438.
Apical whorls 13, sculpture of crowded, minute, oval to
circular pits caused by fusion of radial and spiral riblets,
with a nearly medial keeled ridge that fades out after nuc-
lear whorls. Postnuclear whorls with low, irregular, pro-
tractively sinuated growth wrinkles, occasionally with la-
mellar periostracal extensions. Peripheral keel possessing a
serrated periostracal fringe. Occasional traces of minute,
vague pits arranged in spiral patterns. Microsculpture of
fine, rather widely spaced radial riblets with faint traces
of very fine and crowded spiral riblets. Sutures deep,
channeled, whorls flatly rounded with stronger spiral
Page 258
THE VELIGER
Vol. 12; No. 3
sculpturing yisible, particularly strongly pitted within a-
perture. Color dark olive yellow brown with irregular
lighter streaks. Apex light yellow orange in tone. Umbili-
cus closed by reflection of lip. Aperture very large, subtri-
angular, almost flat above periphery, gently and evenly
rounded below, with very sinuated edge, inclined almost
50° from the shell axis. Height of holotype 2.55 mm,
diameter 5.82 mm.
Holotype: Papua: Arva (= Aroa) River at 6000 feet
elevation. British Museum (Natural History) number
1907.5.28.11.
Range: Known only from the type collection.
Material: Arva River (3 specimens, BMNH 1907.5.28.11,
SMF 161111, ANSP 109258).
Remarks: The much larger Paryphantopsis fultoni has a
flat spire and lacks any trace of postapical spiral puncta-
tions. No other species have a sharp peripheral keel.
Paryphantopsis (Paryphantopsis) latior
VAN BENTHEM JuTTING, 1964
Paryphantopsis latior VAN BENTHEM JuTTING, 1964,
Nova Guinea, Zool., 26: 15 - 16; figs. 20 - 23 — Ok
Minan, beyond bivak 39A at 1450 to 1500m
elevation, Star Mountains, Irian (ca. 140°48’43”
E, 4°54’32”S).
Paryphantopsis (Paryphantopsis) lamelligera
(THIELE, 1928)
Flammulina (Paryphantopsis) lamelligera THir.e,
1928, Zool. Jahrb., Syst., 55: 126; plt. 5, figs. 10, a
— Maeanderberg, 670m elevation, Upper Sepik
River, Sepik District, New Guinea (ca. 147°40’ E,
6°30’S).
Paryphantopsis (Paryphantopsis) striata (Futon, 1902)
Paryphanta striata Futton, 1902, Ann. Mag. Nat.
Hist., (7), 9: 182 — Arva (=Aroa) River, New
Guinea.
Flammulina (Paryphantopsis) striata (Fuuton),
Ture, 1928, Zool. Jahrb., Syst., 55: 126; plt. 5,
fig. 11.
Illonesta striata (FULTON), IREDALE, 1941, Austral.
Zool., 10 (1): 93.
Paryphantopsis (Paryphantopsis) striata (Futon),
SoteM, 1958, Arch. f. Mollusk., 87 (1-3): 23.
Description: Shell very large, with 34 loosely coiled,
globosely swollen whorls. Apex and spire moderately and
evenly elevated, body whorl descending only slightly more
rapidly, H/D ratio 0.707. Apical whorls 13, sculpture of
widely spaced, approximately equal radial and spiral rib-
lets, forming a cross-hatch sculpture except in the sutural
area where the spiral ribbing is absent. Secondary sculp-
ture of very fine radial ribbing, barely visible. Postnuclear
whorls with narrow to low, broadly rounded, vague growth
wrinkles, occasionally prolonged into periostracal lamellae.
Microsculpture of very fine, crowded, radial riblets crossed
by slightly finer spiral riblets. Color reddish brown with
irregular orange-yellow periostracal streaks, apex light
yellow orange. Sutures deep, whorls strongly rounded
above, flattened latcrally above periphery, with flattened
and elongated basal margin. Umbilicus closed. Aperture
very large, somewhat flattened laterally above periphery
and on basal margin, inclined about 30° from the shell
axis, with very sinuate lip edge. Height of holotype
5.69 mm, diameter 8.04 mm.
Holotype: Papua: Arva (—Aroa) River at 6000 feet
elevation. British Museum (Natural History) number
1902.5.28.10.
Range: Known only from the type collection.
Material: Arva River (2 specimens, BMNH 1902.5.28.10,
SMF 161110).
Remarks: The apical sculpture of widely spaced radial
and spiral ribs which form a cross-hatch pattern, is an
obvious forerunner of the minutely pitted sculpture of
Paryphantopsis louisiadarum. In the latter the pattern
of pitting carries over onto the postnuclear whorls, but is
totally absent in the few specimens known of P striata.
The lack of periostracal fringes is shared with P lowisia-
darum, which differs in its larger size, pitted apical sculp-
ture and less rapid body whorl descension.
Paryphantopsis (Paryphantopsis) fultoni (Corn, 1922)
Chronos fultont Corn, 1922, Ann. Mus. Civico Stor.
Natur. Giacomo Doria, (3), 9: 361 - 363; figs. 3-4
— Moroka, New Guinea.
Diagnosis: Shell very large, diameter 7.97 mm to 9.28 mm
(mean 8.52 mm), with 23 to 3 very loosely coiled whorls.
Apex distinctly elevated, spire flat, body whork not des-
cending more rapidly, H/D ratio 0.354 to 0.434 (mean
0.401). Apical whorls 14, bi-keeled, flat above supraperi-
pheral keel, a marked supraperipheral sulcus, followed by
a rounded peripheral keel, sculpture of minute pits ar-
ranged in spiral rows. Postnuclear whorls macroscopically
smooth above periphery except for irregular malleations,
growth wrinkles, and occasional periostracal extensions.
Periphery with very large and irregular periostracal fringe,
—
Vol. 12; No. 3
lower palatal wall with regularly spaced, crowded, nar-
row, prominent periostracal rib extensions. Under 96
magnification traces of micro-radial riblets and finer mi-
crospiral riblets are visible on upper surface, clearly defined
on lower surface of shell. Sutures deep on apex, becoming
very shallow on spire, apical whorls bikeeled, lower whorls
becoming strongly flattened laterally above sharply keeled
periphery, with evenly rounded lower palatal margin.
Columellar lip recurved over and closing very narrow
umbilical chink even in juveniles. Aperture very large,
flattened laterally above periphery, inclined more than
50° from shell axis.
The keeled periphery, very prominent periostracal fring-
ing, absence of spiral pitting below the apex, and large
size identify Paryphantopsis fultoni; P elegans (FULTON)
has the keeled periphery and fringes, but is much smaller
(diameter 5.82mm), has spiral pitting on the lower
whorls, and the sutures deep even to the aperture; P
lamelligera (THIELE) has only a very weak peripheral
keel, much stronger spire and apical elevation with marked
descension of the body whorl. All other Paryphantopsis
have rounded peripheries.
Description: Shell very large, with 2% very loosely coiled
whorls. Apex slightly protruding, spire flat, body whorl
not descending, H/D ratio 0.424. Apical whorls slightly
less than 14, flattened above mid-upper palatal keel, con-
cave below to peripheral keel, sculpture of minute punc-
tations formed by fusion of spiral and radial ribs. Post-
nuclear whorls with irregular growth wrinkles, sometimes
bearing periostracal fringe extensions, surface somewhat
malleated with remnants of apical keels and sulci, peri-
phery with roughly triangular, quite large periostracal
fringes, lower palatal wall with narrow, high, rather
crowded, regularly spaced radial periostracal rib exten-
sions. Microsculpture visible on lower surface as fine
radials crossed by finer spirals, barely visible on upper
surface. Sutures deep on apex, becoming quite shallow
on lower spire, postnuclear whorls flattened laterally above
sharply angled periphery, evenly rounded below. Color
reddish brown near aperture, becoming greenish yellow
on upper spire, apex yellow-white. Umbilical chink closed
by reflection and periostracal extension of columellar lip.
Aperture very large, flattened laterally above periphery,
inclined about 50° from shell axis. Height of lectotype
3.46 mm, diameter 8.11 mm.
Lectotype: Moroka, Laloki River, headwaters, east of
Port Moresby. Collected by Lamberto Loria in July 1893.
Museo Civico di Storia Naturale “Giacomo Doria,”
Genova.
Material: Moroka (19 specimens, MHNG, FMNH
159265).
THE VELIGER
Page 259
Remarks: Differences from Paryphantopsis elegans are
covered in the diagnosis above. Probably the most signifi-
cant are the absence of spiral sculpture on the lower
whorls and regular fringed sculpture on the shell base
seen in P fultoni. Moroka is a classic ornithological col-
lecting locality on the slopes of Mt. Wori-Wori about 25
miles inland of Tupuseleia and fairly near Port Moresby.
Paryphantopsis (Paryphantopsis) louisiadarum
(MG6LLENDorr®rF, 1899)
Paryphanta louisiadarum MO.LLENDoRFF, 1899,
Nachr.-Bl. dtsch. Malak. Gesell., 31: 89 — Louisiade
Islands, New Guinea; MOLLENDorFF, 1902, Syst.
Conch: ‘Cab:, I-12; B, p: 17; pit. 3 figs: 1-3;
SoteM, 1959, Arch. f. Mollusk., 88 (4-6) : 156; plt.
12, figs. 10, 11; plt. 13, fig. 6.
Illonesta louisiadarum (MO6OLLENDORFF), IREDALE,
1941, Austral. Zool., 10 (1): 93.
Paryphantopsis (Paryphantopsis) louisiadarum
(MOLLENDoRFF), SoLem, 1958, Arch. f. Mollusk.,
87 (1-3): 23 — Rossel Island, Louisiades.
Description: Shell very large, with 2% very loosely coiled,
globosely swollen whorls. Apex nearly flat, lower whorls
descending moderately, H/D ratio 0.704. Apical whorls
13, sculpture a network of spiral and radial ribs broadly
joining each other, forming regular rows of small circular
to oval pits, about 15 in number across the whorl. Pitted
sculpture continuing on post nuclear whorls with radial
ribbing gradually becoming more prominent and at the
end of the second whorl a fine microsculpture of radial
riblets crossed by much finer and more crowded spiral rib-
lets becoming visible. On the body whorl the sculpture has
become reduced to series of spiral grooves in which the
small pits are visible with most of the surface showing
irregular growth lines and microsculpture. Color olive
yellow brown with occasional darker flammulations be-
coming very dark in the sutures. Umbilicus closed by
reflection of lip. Aperture very large, somewhat flattened
laterally above periphery and on basal margin, inclined
about 35° from the shell axis. Height of lectotype 6.54
mm, diameter 9.28 mm.
Lectotype: New Guinea: Louisiade Islands. Natur-Mu-
seum Senckenberg, Frankfurt number 137274.
Range: Louisiade Islands.
Material: Rossell Island (1 specimen, ANSP 109257) ;
Louisiades (2 specimens, SMF 137274, SMF 165564).
Remarks: Only the sharply keeled Paryphantopsis fultoni
approaches the size of this species. No anatomical material
Page 260
is available and apparently no specimens have been col-
lected in this century.
Paryphantopsis (Gallodema) globosa (HEDLEY, 1890)
Rhytida globosa Hepiey, 1890, Ann. Rep. British
New Guinea, 1888 - 89: 65 — Mt. Victoria, Owen
Stanley Mts., Papua; Hepiey, 1891, Proc. Linn.
Soc. New South Wales, (2), 6: 80; plt. 10, figs. 15,
16; Sorem, 1959, Arch. f. Mollusk. 88 (4-6): 156
tomlaye
Gallodema globosa (Hepiry), IREpALE, 1941,
Austral. Zool., 10 (1): 92.
Paryphantopsis (Gallodema) globosa (HEDLEY),
Sotem, 1958, Arch. f. Mollusk., 87 (1-3): 24
Remarks: I have not been able to examine the type of
Rhytida globosa, which was deposited in the Queensland
Museum, Brisbane. The original description mentioned
the pitted apical sculpture and “bleached” early whorls.
A single shell from Moroka (MHNG) collected by Lam-
berto Loria in July, 1893, is referred to this species, al-
though much smaller in size. It is 4.44 mm high, 5.95 mm
in diameter, H/D ratio 0.747, with 33 whorls. HEDLEy’s
original measurements of height 10 mm and diameter 17
mm give a calculated H/D ratio of 0.588, but the original
figures give a height of 10.9mm, diameter 15.9 mm for
a calculated H/D ratio of 0.686. I consider that the Moro-
ka shell is closely related to HEDLEY’s species, if not merely
a juvenile example.
Apical sculpture of the Moroka shell consists of co-
equal radial and spiral riblets that produce a lattice pat-
tern with a few worn spots appearing “pitted” as in
typical Paryphantopsis. IREDALE (loc. cit.) sketchily de-
scribed a new genus, Gallodema, for this species. It is
intermediate between Pilsbrycharopa and Paryphantopsis
in shape and whorl count. The apical sculpture also ap-
pears transitional. I question the desirability of generic
separation and prefer to use Gallodema as a subgenus of
Paryphantopsis.
COMPARATIVE REMARKS
Despite knowing only part of the visceral hump anatomy
for one species in each genus, a comparison between the
two yields significant information on patterns of pulmonate
evolution. Pilsbrycharopa and Paryphantopsis belong to
the same subfamily. They have quite different average
patterns of shell structure. The two dissected species,
Pilsbrycharopa gressitti and Paryphantopsis dauloensis,
THE VELIGER
Vol. 12; No. 3
are sympatric. Therefore existing genital differences may
provide no information helpful in elucidating phylogeny.
In the Pacific Island endodontid taxa, sympatric species
in the same genus or closely related genera show marked
character displacement in penial structures (unpublished
data). One species of such a pair or group may be
strikingly altered in penis structures, while the other(s)
may be only very slightly modified, or they may diverge in
different directions from the “‘average” structural pattern.
I cannot state whether the Pilsbrycharopa or the Pary-
phantopsis is greatly modified, since without dissection of
many specics, determination of the average pattern is not
possible. Pilsbrycharopa gressitti has an oval penis, lacks
a vergic papilla, has a large stimulatory pad and large
smooth pilasters near the base, while Paryphantopsis dau-
loensis has a club-shaped penis with a vergic papilla, no
stimulatory pad, and corrugated longitudinal pilasters. The
existence of these differences may have phylogenetic sig-
nificance or may reflect character displacement to enforce
species isolation. A hint of possible ecological divergence
is given by the body color, yellow-white in Paryphantopsis,
greyish tones in Pilsbrycharopa. In the Pacific Island en-
dodontids, yellow-white body color is associated with litter
dwelling forms, greyish tones with semiarboreal taxa. No
ecological data have been recorded concerning the habitat
of either genus, but general affinities of both seem to be
with the Pacific Island, rather than the New Zealand-
Australian taxa. Correlation of ecological zonation and
body color reasonably could be expected to hold for the
New Guinea taxa. A combination of slight ecological di-
vergence and exaggeration of species isolating mechanisms
is the common pattern when closely related species become
sympatric. A possible hint of dietary difference is given
in the radular teeth. Those of Paryphantopsis have the
basal plates of the laterals much narrower and the mar-
ginal basal plates are square rather than rectangular.
Mesoconal cusps are shorter in Paryphantopsis and there
is no evidence of ectoconal cusp splitting on the outermost
marginals. This shift in radular tooth shape and cusp size
may indicate a scraping feeding action rather than the
normal slicing action in more typical endodontid radulae.
Ecological and life history observations in the Daulo Pass
area on the two genera could yield very important data
on the origin and maintenance of mating barriers in
pulmonates. Both genera also have been collected at
Moroka.
While the above data concerning such factors of micro-
evolution are interesting, differences between the two gen-
era provide deeper insights into patterns of major change.
A look at the shell illustrations of the two new species
(Figure 1) and mean measurements (Tables 2, 3, 5) con-
firms the existence of several consistent differences in
Vol. 12; No. 3 THE VELIGER Page 261
Gf?
ila
Gy E
® ®)
Figure 3
Visceral hump volume relationships and foot length-pallial cavity
ratios in Pilsbrycharopa (a) and Paryphantopsis (b)
Page 262
coiling pattern, shape and sculpture. Table 6 summarizes
the major contrasting shell features between the two gen-
era. There is some overlap in umbilical width and apical
sculpture, plus considerable overlap in diameter, but the
basic contrast in structure and form is obvious. Within
the context of the Endodontidae, the characteristics of
Paryphantopsis represent much greater departures from
the typical structural pattern. “Cope’s Law” concerning
the general tendency towards size increase during a phylo-
genetic series is consistent with the greater size of Pary-
phantopsis.
Within the subfamily Charopinae, spiral apical cording
is primitive. Addition of radial elements clearly is second-
ary. The increase in radial apical sculptural elements in
the species group of Pilsbrycharopa is correlated with
larger size. The peculiar “pitted” apical sculpture of Pary-
phantopsis (sce SoLeM, 1959, plt. 13, fig. 6) is most easily
derived from partial height reduction of a radial and
spiral rib network. It is a secondary derivation after
radial clements are large enough to equal the spiral
cording. Paryphantopsis is much more specialized in apical
sculpture.
The most significant alteration is the reduction in whorl
count and correlated change in coiling pattern seen in
Paryphantopsis. While accurate area measurements could
not be made, casual inspection shows that the apertural
cross-sectional area is much less in the 4- to 5-whorled
Pilsbrycharopa (Figure 1, e) than the 3-whorled Pary-
phantopsis (Figure 1, b). It is equally obvious that, in
shells of equal diameter, lowering the whorl count will
significantly lower the total linear distance from the lip
edge to shell apex as measured along either the periphery
or midwhorl line. Using the illustrations of Pilsbrycharopa
gressitti and Paryphantopsis dauloensis as examples and
adjusting for the size differences, the distance in the
Paryphantopsis is only 73% of the measured midwhorl
distance in the Pilsbrycharopa. The shell serves to encom-
pass the visceral hump and provides space for withdrawal
of the head, foot and tail, so that alterations in shell form
cannot be segregated from a consideration of changes in
the anatomy. Actually, since the shell is deposited by the
mantle edge of the living snail, it is fallacious to think
of the two separately. Practically speaking, however, chan-
ges in the shell form are more easily perceived and docu-
mented so that an initial explanation of the changing
surface cover (=shell) of the visceral hump permits
better understanding changes in organs of the latter.
The net effect of a reduction in whorl count and loosen-
ing of the coiling patterns is shown in Figure 3. If the two
visceral humps could be decoiled and distorted into simple
cones, the volume would change only slightly, but the
spire angle would be enlarged, the height of the cone
THE VELIGER
Vol. 12; No. 3
shortened, and the aperture widened in Paryphantopsis.
There is no doubt that Paryphantopsis has a shorter, wider
area for the visceral hump organs. Since this pattern of
reduction in whorl count and loosening of coiling pattern
is repeated in family after family of land snails and is an
initial step in the process by which snails evolve into
slugs, examination of the anatomical changes in the vis-
ceral hump region between Pilsbrycharopa and Paryphan-
topsis is worthwhile. When combined with similar studies
in other taxa, such work may provide criteria for evalu-
ating the affinities of slugs.
In this report I wish to focus on two organ complexes,
the entire pallial region and the genitalia. Unfortunately
I could not examine the apical genitalia of either genus,
but changes in this area are fairly standard.
Pallial regions of both genera are illustrated in Figures
2a and 2f. Since the mean diameter of Pilsbrycharopa
gressittt is 6.76 mm and the mean diameter of Paryphan-
topsis dauloensis is 5.48 mm, or 19% less, direct measure-
ment comparisons are less meaningful than the use of
ratios. Total length of the pallial cavity in Pilsbrycharopa
is 6.25 mm (0.92 shell diameter); 2.96 mm (0.54 shell dia-
meter) in Paryphantopsis. While in Pilsbrycharopa it
occupies more than one-third of the body whorl, in Pary-
phantopsis it is reduced to about one-sixth, or only half
the proportionate amount. No accurate measure of pallial
cavity width was possible, since the delicate roof tissue on
the lower palatal wall is inevitably torn during dissection
and further cut in pinning the cavity out for detailed
study. It undoubtedly is greater, but the organs all occu-
py the upper palatal portion and parietal-palatal margin,
so that this dimension can be ignored.
In most land snails with normally developed shells the
hindgut follows the parietal-palatal angle from the anus
past the pallial cavity apex. In the Charopinae, the
kidney is bilobed, with a rectal lobe lying alongside the
hindgut from anterior end to the curved base of the kid-
ney and a pericardial lobe extending anterior of and
posterior to the heart. Pilsbrycharopa departs from this
pattern in that the kidney is partially rotated downwards,
with only the anterior third of the rectal kidney arm in
contact with the hindgut. This is carried further in Pary-
phantopsis, where only the anterior eighth of the rectal
kidney arm is along the hindgut. That this is *basically
simple rotational change is shown by calculating the
width to length ratios for the kidneys, which are 0.69 in
Paryphantopsis and 0.65 in Pilsbrycharopa. The difference
is within the probable range of measurement error and
has no significance. Pilsbrycharopa represents an initial
stage in pallial cavity shortening, which is carried further
in Paryphantopsis. A more obvious change concerns the
spatial relationships of the primary (ascending) and sec-
Vol. 12; No. 3
THE VELIGER
Page 263
ondary (descending) ureters (KD). In typical Charopi-
nae the two branches are either compacted together
between the kidney lobes or a narrow strip of lung roof
is visible between and a very slight angle is developed.
In Pilsbrycharopa the ureter arms are clearly divergent
with about a 28° angle. In Paryphantopsis the angle is
greatly increased, reaching about 60° in dissected speci-
mens. Obviously, continuation of this trend would result
in development of a primary ureter transversely oriented
in the pallial cavity, essentially paralleling the mantle
collar edge. Such an arrangement is present in the Suc-
cineidae. The phylogenetic implications of this will be
considered in another report.
Two additional pallial alterations require comment. In
Pilsbrycharopa the external ureteric pore (KX) is located
distinctly behind the anal opening (A) ; in Paryphantop-
‘sis the two open side by side at the posterior pneumostomal
margin. Pilsbrycharopa has the principal pulmonary vein
(HV) simple and unbranched; Paryphantopsis has mul-
tiple branches on the pallial roof.
From a pallial cavity extending one-half to five-eighths
of a whorl apically in most Charopinae, Pilsbrycharopa
shows a reduction to one-third of a whorl and Paryphan-
. topsis to one-sixth. The proportionate shortening of the
pallial cavity reduced the length of the pulmonary vein
and hence the surface for both water and gas exchange.
Branching of the vein restores balance to the system. Ad-
justment of the kidney and ureter to the “squeeze” was
achieved by partial rotation of the kidney and opening
of the angle between the primary and secondary ureter.
This is only one of the many ways to compensate for the
changed forms. The species described as Flammulina
nigrescens VON MOLLENDoRFF (1900, pp. 107-109) from
Ponape, Caroline Islands has a shell form very similar to
that of Paryphantopsis, but the pallial region is very dif-
ferent. In that species, the very elongated, bilobed kidney
has become square in shape, very thick in the middle, the
ureter arms right next to each, and the kidney does not
deflect from the hindgut (unpublished data).
Genital structures of the two genera agree in essentials
and differ widely in details. I have dissected over 100
species of endodontids from the Pacific Islands. Pilsbry-
charopa and Paryphantopsis differ sharply from all of these
in having a diverticulum (EL) on the epiphallus and in
having the penial retractor muscle insert on the epiphallus
rather than on the head of the penis. Structures of the
prostate-uterus, spermatheca and vas deferens agree with
the typical Charopinae. The penial differences were out-
lined above and are not discussed further. The most ob-
.vious difference in the two genera is the length of the
penial retractor muscle (PR) — long in Pilsbrycharopa,
very short in Paryphantopsis. Since the muscle arises from
the diaphragm near the pallial cavity apex, shortening of
the cavity shortens the muscle.
In Paryphantopsis the epiphallus and vagina are con-
siderably longer than in Pilsbrycharopa. At first glance
this appears anomalous, since shortening of the visceral
hump in the former genus should tend to shorten the
genital organs. Unfortunately the complete prostate-uter-
ine areas of neither genus were available for study. From
dissections of Pacific Island and Thailand taxa, I have
learned that shortening of genital organs does not proceed
uniformly, but is done on a zonal basis. Hence shortening
may be restricted to the prostate-uterine section in this
particular Paryphantopsis, but have proceeded in the
lower area in the Pilsbrycharopa. The extent and varia-
bility of such zonal compaction can be appreciated by
comparing the genital anatomy of Durgella libas and
Cryptaustenia gadinodromica (see SoteM, 1966, p. 54,
fig. 9a; p. 62, fig. 12b). Both have shells of about 4 whorls
and very large and elongated feet. In Durgella (loc. cit.,
p. 54) the prostate-uterus is very long and the free oviduct,
spermatheca and penis obviously short; in Cryptaustenia,
the prostate and uterus are only slightly longer than the
spermatheca and penial complex. Megaustenia siamensis
(Ibid., p. 83, fig. 19a) agrees more with Durgella, but has
the penial complex convoluted and coiled. All three
helicarionid genera represent about 14 whorls reduction
from the normal helicarionid pattern. Compaction of
genital structures has occurred, but different zones have
been affected to different extents.
This makes it exceedingly difficult to evaluate differ-
ences betwen the genitalia of two species that show par-
tial visceral hump reduction, unless considerable data are
available concerning the “normal” or “‘typical” structural
pattern in species that do not show such shell reduction.
Since Pilsbrycharopa and Paryphantopsis show major dif-
ferences from Pacific Island genera, 7. ¢., possession of the
epiphallic diverticulum, insertion of the penial retractor
on the epiphallus, adjustment of the pallial cavity to com-
paction by kidney rotation, and internal penial configura-
tions, we do not know what the typical pattern is in
related genera.
SUMMARY
Pilsbrycharopa and Paryphantopsis present contrasting
patterns of shell structure (Table 6), although they are
very similar in anatomy. Changes from the normal charo-
pinine pattern in the pallial complex of these genera indi-
cate one method of coping with an early stage in visceral
Page 264
THE VELIGER
Vol. 12; No. 3
Table 6
Shell Differences
Between Pilsbrycharopa and Paryphantopsis
Pilsbrycharopa Paryphantopsis
Whorl count 33 to5+ 23 to 34
Coiling pattern tighter much looser
Umbilicus usually widely slight lateral crack
or moderately or completely
open closed
Size of aperture smaller much larger
reduced to
periostracal fringes
spiral and radial
Radial sculpture usually typical
Apical sculpture spiral cords to
equal radials to pitted
and spirals
Calcification average to below average
heavy to slight
Diameter 1.9-7.3mm 3.6 - 17.0mm
median 3.88 mm | median 5.8 mm
hump reduction. If carried to a greater extent, evolution
of slug-like forms becomes possible and interpretation of
such patterns will aid in determining the affinities of slugs.
Pilsbrycharopa contains nine species. When more species
can be dissected, it may be broken up into several genera,
but in the absence of such data, I prefer to use a single
broadly defined generic taxon. Several of the species may
be compound taxa.
Paryphantopsis contains fourteen nominate units. They
are known only from type material and the extent of intra-
and inter-populational variation is unknown.
LITERATURE CITED
ARcHBOLD, Ricuarp, A. L. Rano «& L. J. Brass
1942. _—_ Results of the Archbold Expeditions. No. 41. Summary
of the 1938-1939 New Guinea Expedition. Bull. Am. Mus.
Nat. Hist. 79 (3): 197 - 288; 3 maps; plts. 1 - 35
BortrTcer, CAESAR R.
1908. Zur Fauna von Amboina (Mollukken).
deutsch. Malakol. Gesell. 40 (4): 180 - 192; 6 figs.
Corn, Giorcio Sitvio
1922. Nuove descrizione di specie di molluschi del Museo
Civico di Genova. Ann. Mus. Civ. Genoa 9 (3) : 359 - 363;
Nachr. Bl.
4 figs.
Franc, A.
1952. | Remarques suivies de rectifications de nomenclature sur
des coquilles Néo-Calédoniennes. Bull. Soc. Zool. France
77: 76-79
Futon, H.
1902. Descriptions of new species of Land-Mollusca from New
Guinea. Ann. Mag. Nat. Hist. 9 (7): 182 - 184
HEDLEY, CHARLES
1890. Description of a new Rhytida from New Guinea.
Ann. Rep. British New Guinea, 1888-1889: 65
1891. The land-molluscan fauna of British New Guinea.
Proc. Lin. Soc. N. S. W, 6 (2): 67-116; plts, 9- 12
IREDALE, TOM
1941. A basic list of the land Mollusca of Papua.
Zool. 10 (1): 51-94; plts. 3 - 4.
MOLLENDoRFF, O.
1892. Die Landschneckenfauna der Tenimber-Inseln (Timor-
laut). Nachr. Bl. deutsch. Malakol. Gesell. 24 (5-6): 81 - 120
plt. 1
Austral,
1899. Neue Arten aus der Strubell’schen Sammlung. _Nachr.
Bl. deutsch. Malakol. Gesell. 31 (5): 89-92
1900. The land shells of the Caroline Islands. Journ,
Malac. 7 (5): 101 - 126; 5 charts; 3 figs.
MOo.venporrr, O. «& W. KoBELT
1902-1905. Die Raublungenschnecken (Agnatha). Erste Ab-
theilung: Rhytididae & Enneidae. Syst. Conch. Cab. I, 12, B;
1 - 362; plts. 1-41
RENscH, BERNHARD
1930. | Neue Land-Pulmonaten von den Kleinen Sunda-Inseln.
Zool. Anz. 89 (3/4): 73-88; 16 figs.
1932. Die Molluskenfauna der Kleinen Sunda-Inseln Bali,
Lombok, Sumbawa, Flores, und Sumba. II. Zool, Jahrb.
Syst. 63: 130 pp.; 3 plts.; 56 figs.
1935. Zur Landschneckenfauna von Timor. Sitz. Gesell.
Naturf. Freunde Berlin 1935: 311 - 336; 19 figs,
RenscuH, I.
1937. | Systematische und tiergeographische Untersuchungen
uber die Landschneckenfauna des Bismarck-Archipels. II.
Arch. f. Naturgesch., N. F 6 (4): 526 - 644; 54 figs.
SmirH, Epcar ALBERT
1896. | On some land shells from New Guinea and other neigh-
boring islands, with descriptions of new species. Journ.
Malacol. 5 (2): 17-22; plt. 2
SoLtem, ALAN
1958. | Endodontide Landschnecken von Indonesien und Neu
Guinea. Arch. Molluskenk. 87 (1-3): 19-26; 6 figs.
1959. On the family position of some Palau, New Guinea,
and Queensland land snails. Arch. f. Molluskenk. 88 (4-6):
151 - 158; plts. 12, 13; 2 figs.
1964. A collection of non-marine mollusks from Sabah.
Sabah Soc. Journ. II (1-2): 40 pp.; 5 figs.
1966. Some non-marine mollusks from Thailand, with notes on
classification of the Helicarionidae.
24: 110 pp.; 3 plts.
Soos, L.
1911. On a collection of land shells from New Guinea and
adjacent Islands. | Ann. Mus. Hungar. IX: 345 - 356; 10 figs.
Spolia Zool. Mus. Haun.
THIELE, JOHANNES
1928. Mollusken vom Bismarck-Archipel, von Neu-Guinea
und Nachbar-Inseln. Zool. Jahrb. 55: 119 - 146; 1 fig.; plt. 5
VAN BENTHEM JurtiNc, W.S. S.
1964. Non-marine molluscs of W. New Guinea. Part 3, Pul-
monata, I. Nova Guinea, n.s. 10 (26): 1-74; 1 map; 62
figs.; plts. 1, 2
Vol. 12; No. 3.
THE VELIGER
Page 265
Reproductive Cycle of the Coot Clam, Mulinia lateralis (Say),
in Long Island Sound *?
BY
ANTHONY CALABRESE
Bureau of Commercial Fisheries, Biological Laboratory, Milford, Connecticut 06460
(Plates 37, 38; 1 Text figure)
INTRODUCTION
THE cooT cLAM, Mulinia lateralis (Say, 1822), a member
of the family Mactridae, has received very little attention
in spite of its abundance in favorable environments. This
ecologically significant clam is a food of many bottom-
dwelling and bottom-feeding animals, including black
drum, Pogonias cromis (BREUER, 1957); scup, Stenotomus
chrysops, other fishes (VERRILL, 1873) ; starfish, Asterias
forbesi (Drsor, 1848), oyster drills, Eupleura caudata
and Urosalpinx cinerea (Say, 1822) (C. L. MacKenzie,
Jr., personal communication) ; and the greater scaup duck,
Aythya marila, and lesser scaup duck, Aythya affinis
(Cronan, 1957).
Periodic histological examination of gonad tissues during
several successive years has been valuable for determining
the periodicity of gametogenesis in many marine inverte-
brates (Grese, 1959). The reproductive cycle of many
species of pelecypods has been described, but until Ropes
(1968a) discussed the reproductive cycle of the surf clam,
Spisula solidissima, gametogenesis had not been described
for any of the Mactridae.
The spawning season for Mulinia lateralis, as indicated
by the presence of larvae in plankton samples, has been
reported to be from mid-July to early September at Prince
Edward Island, Canada (Sutuivan, 1948). Loosanorr,
Davis & CHANLEy (1966) reported these larvae to be
extremely numerous in plankton samples from Long Is-
land Sound during late summer, but they did not attempt
to define the spawning season. SHaw (1965) determined
' Part of a dissertation submitted to the Graduate Faculty of the
University of Connecticut in partial fulfillment of the require-
ments for the degree of Doctor of Philosophy.
2 Contribution No. 58 from the Marine Research Laboratory,
University of Connecticut.
from collections with Thorson bottles that M. lateralis
spawns and sets in the Tred Avon River, Maryland, from
May to November; peak setting is in September (Hanks,
1968).
Knowledge of the reproductive cycle of this clam is
essential to an understanding of larval production and,
ultimately, to the abundance of this ecologically impor-
tant bivalve. The duration of the spawning season of Mu-
linia lateralis and the time of appearance of the larvae
were determined by following gametogenic development
in histological sections throughout the year and by making
plankton studies during the spawning season.
MATERIALS ann METHODS
Mulinia lateralis were collected from several areas in the
Bridgeport-Milford-New Haven, Connecticut, area of
Long Island Sound. They were kept in boxes of sand
placed on an underwater dock in Milford Harbor or in
outdoor running water tanks at the laboratory and main-
tained as specimens to be sacrificed for histological study.
Ten clams were collected from this supply either weekly
or in alternate weeks from August 2, 1965 to August 28,
1967, and placed in Lillie’s decalcifier fixative (HuMa-
son, 1962) for 24 to 48 hours. The gonad was dissected
out, dehydrated in alcohol, cleared in xylene, and embed-
ded in paraffin by standard techniques. Gonad tissues were
sectioned at 74 with a rotary microtome, stained with
Delafield’s hematoxylin, and counterstained with eosin.
The sections were examined under an AO Spencer light
microscope at X 100 and Xx 430 magnification; the go-
nad tissue was assigned to one of the stages of development
described by Ropes & STICKNEY (1965), who categorized
the seasonal cycle of gametogenesis in the soft-shell clam,
Mya arenaria LINNAEUS, 1758, as follows: inactive, ac-
Page 266
tive, ripe, partially spawned, and spent. Photomicrographs
of the various stages of gametogenesis were taken by use
of a standard Zeiss light microscope and camera at X 200
and < 500 magnification.
To provide corroborative evidence on the beginning and
duration of the spawning season of Mulinia lateralis, plank-
ton samples were collected twice weekly at 5 stations in
the Bridgeport-Milford-New Haven area from June 17 to
October 3 and on October 8 and 24, 1968. By examining
the samples I was able to determine when the larvae first
appeared and to follow variations in their abundance.
Samples were collected with the research vessel, Shang
Wheeler, by pumping a 200-gallon water sample through
a no. 10 bolting-silk net to screen off the larger, more
easily identifiable larvae, and then through a no. 20 net
to collect smaller larvae. A 12 2-inch iron suction head
was lowered to mid-depth with a 1.5-inch rubber hose to
collect samples. The opening of the suction head was
covered with fine-mesh copper screening to prevent en-
trance of extraneous debris. Two metal drums connected
in tandem provided a system whereby water flowing
through the first net was trapped in the first drum and
passed through the second net through an overflow pipe.
The samples were washed into 150-ml jars containing 1
ml of formalin as a preservative and later examined mi-
croscopically for the presence of M. lateralis larvae.
HISTOLOGICAL STUDY
oF GAMETOGENESIS
The gonad in ripe Mulinia lateralis is a large and clearly
defined organ and easily distinguishes ripe individuals
from those with undeveloped gonads or those that have
discharged all or most of their gametes. The gonad of
ripe animals is an almost uniform, continuous mass of tis-
sue surrounding the digestive tract and the digestive di-
verticula. During the spawning period the thickness of the
gonad decreases as a result of the discharge of gametes.
The gonad of spent animals is scarcely discernible by visual
examination, but can be observed readily in histological
sections.
The data from two years of monthly sampling for
histological study were combined since progress of game-
togenesis was essentially the same both years (Text figure
THE VELIGER
Vol. 12; No. 3
Inactive Gametogenesis
Ei Active Gametogenesis
Partially Spawned
BH Ripe
[_] Spent
Ea a se
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Month
Figure 1
Gonad condition of Mulinia lateralis in Long Island Sound
from August 1965 to August 1967. The length of each shaded area
represents the percentage frequency of clams in each category.
The numbers above each bar represent the total number of clams
sampled that month. Data from the two years of observation are
combined.
1). Histological examination of these samples indicates
that spawning of Mulinia lateralis is completed by the end
of September or early in October. No ripe unspawned indi-
viduals were found in October; 39.5% were partially
spawned and 36.8% were completely spent. The gonads
of spent clams examined microscopically during this pe-
riod showed large, distended follicles containing few or no
spermatozoa or ova within the lumina (Plate 37, Figure
5; Plate 38, Figure 11), whereas the follicles of partially
spawned animals contained small numbers of ripe gametes
(Plate 37, Figures 3, 4; Plate 38, Figure 10).
Sections of gonads fixed in late July and August re-
vealed that several animals were beginning to undergo
active gametogenesis. Whether this was preliminary to a
second annual spawning was not determined. The gonad
follicles at this time contained many spermatogenic and
oogenic cells in various stages of development (Plate 37,
Figure 1; Plate 38, Figure 7). The percentage of clams
showing gametogenesis increased through November even
though the water temperature decreased to 6° C, and by
December about 80% of the animals examined: were un-
dergoing gametogenesis (Text figure 1). This activity con-
tinued at an increased level through January, February,
Explanation of Plate 37
Section of Gonad Tissue of Male Mulinia lateralis
Figure 1: Active phase of spermatogenesis.
Figure 2: Fully mature. (XX 500)
Figure 3: Partially spawned. (XX 200)
(X 500)
Figure 4: Partially spawned. (XX 500)
Figure 5: Spent, with few sperm retained.
Figure 6: Inactive phase. (XX 500)
(X 200)
Tue VELIcER, Vol. 12, No. 3 [CALABRESE] Plate 37
Wa : pee
; ait
re « steed ! : «
Figure 3
Vol. 12; No. 3
and March, and by April the gametes in 1 of 38 animals
sectioned were morphologically ripe.
Experiments with Mulinia lateralis made it evident that
at least some gametes of this species were both morpho-
logically and physiologically ripe at times other than
during the normal spawning period. LoosanorF (1937)
reported the possibility that the hard-shell clam, Merce-
naria mercenaria (LINNAEUS, 1758), contained physio-
logically and morphologically ripe gametes at times other
than during the immediate prespawning or spawning pe-
riod. In one experiment with Mulinia lateralis, animals
that were collected on January 10 at a water temperature
of -0.1° C were brought into the laboratory and both
males and females were induced to spawn immediately
when placed in water of 20° C. The eggs released were
fertilizable, but larval development was poor. In another
experiment, animals collected on January 23 at a water
temperature of 4° C were immediately induced to spawn
in the laboratory. They released eggs that developed into
normal straight-hinge larvae within 24 hours after fertil-
ization.
By mid-April the water temperature in the outdoor
tanks increased to about 7° C; gametogenic activity also
increased and one ripe female was found. As stated pre-
viously, however, some animals had follicles with at least
a few morphologically ripe gametes throughout the win-
ter. As the water temperature increased to about 10°C
in mid-May, vigorous gametogenesis was resumed and
38.3% of the adult animals examined were ripe. This
rapid proliferation and maturation of sex cells continued
through June and July. Follicles of ripe males were
crowded with spermatozoa whose tails filled the center of
the lumina (Plate 37, Figure 2). Each follicle of ripe
females contained many mature ova which appeared to be
free in the lumen or attached to the follicle walls by
slender stalks (Plate 38, Figures 8, 9).
The first partially spawned individuals were in samples
taken July 10 to 12, when the water temperature had
reached approximately 20° C (Text figure 1). By the end
of August 61.3% of the animals examined were partially
spawned and a few were spent. Slight gametogenic activi-
ty began again in late July and August, followed by rapid
proliferation of sex cells in September. More ripe clams
were found in September than in August, but it was not
determined whether this gametogenic activity produced
a second wave of spawning. During the colder months,
from October through April, a few individuals appeared
to be in the inactive stage (Plate 37, Figure 6; Plate 38,
Figure 12). At this stage it was most difficult to distinguish
between the sexes. The presence of one or two ova or
spermatozoa in some follicles, however, enabled me to sex
_ the animals.
THE VELIGER
Page 267
Of a total of 597 Mulinia lateralis examined, 302 (50.6
per cent) were females and 295 (49.4 per cent) were
males. No hermaphrodites were found. Hermaphroditism
is rare in the closely related surf clam, Spisula solidissima,
and then is apparently a developmental accident (RopEs,
1966, 1968b).
PLANKTON STUDIES
It was possible to study the occurrence of Mulinia lateralis
larvae in the plankton only during the summer of 1968
when about 350 samples were collected and examined.
Larvae first appeared in the plankton on July 8, when the
water temperature ranged from 16° to 20° C at the vari-
ous stations sampled. Since it was impossible to identify
positively all bivalve larvae in a sample, exact numbers of
M. lateralis larvae were not recorded. However, 2000 to
3000 M. lateralis larvae per 200-gallon sample were con-
sidered abundant and 10 to 20 larvae considered few.’
Larvae were abundant on July 23 at water temperatures
of 19° to 21° C. Numbers were reduced drastically by July
29 and no larvae were found in samples collected August
1 to 8. Larvae were again present on August 12 and were
abundant in samples of August 19 to 22. The number of
larvae decreased again in late August and by September
3 only a few were in the plankton samples. Numbers con-
tinued to be low through October 8, but even on October
24 (when the last samples were collected) the samples
contained a few larvae. Water temperature by this time
had decreased to about 17° C.
MODE or REPRODUCTION
AND PELAGIC EXISTENCE
The examination of gonad sections suggests several gen-
eralizations about the reproductive cycle of Mulinia la-
teralis. Gametogenesis is essentially continuous throughout
the year, i.e., there is no completely inactive period in
winter or summer. Some individuals at each sampling pe-
riod showed gametogenic activity, but gametogenesis was
most active and more ripe gametes were present from mid-
July through August. A spawning peak was reached in
August and development of ripe cells again increased in
September. It was not determined whether this September
development indicates a second reproductive cycle because
gametogenesis does not totally cease before this period.
The abundance of larvae in plankton samples disclosed
that two peaks of spawning occur; one is in late July, just
after spawning begins (as determined from histological
sections), and the second and greater one is in middle to
Page 268
THE VELIGER
Vol. 12; No. 3
late August, which coincides with the increased percent-
age of partially spawned animals in my histological prep-
arations (Text figure 1). This second peak of spawning
appears to be the major one for the normal reproductive
period, as noted by LoosanorF et al. (1966), who stated
that Mulinia lateralis larvae were extremely numerous in
plankton samples from Long Island Sound during the lat-
ter part of the summer. Sutuvan (1948) found M.
lateralis larvae from mid-July to September in plankton
samples from Malpeque Bay, Prince Edward Island, Can-
ada, a period comparable to that in the present study.
Even though Malpeque Bay and Long Island Sound are
widely separated geographically, their temperature re-
gimes are similar. SULLIVAN first observed M. lateralis
larvae in plankton samples about July 18, when the water
temperature was about 21° C, and continued to take lar-
vae until early September, after which time the water
temperature had decreased to about 17°C. These dates
and temperatures correspond closely with the data of the
present study. This similarity tends to indicate that these
two populations of M. lateralis, even though widely sep-
arated geographically, are physiologically similar. The
temperature requirements for gonad development and
spawning of both groups appear to be the same.
SHaw (1965) collected recently set Mulinia lateralis |
larvae in modified Thorson bottle collectors in the Tred
Avon River, Maryland, from May into November and re-
ported that this clam appeared to have a continuous peri-
od of setting with no apparent gaps during the spawning
season. Although he did not report temperature data for
this period, I determined from other sources that water
temperatures in the Tred Avon River from May to No-
vember approximated those in Long Island Sound from
July to September. It appears, therefore, that M. lateralis
from Long Island Sound and the Tred Avon River, Mary-
land, may have similar temperature requirements for
gonad development and spawning.
Ropes & STICKNEY (1965) reviewed the information
available on the reproductive cycle of Mya arenaria and
indicated that two important phenomena occurred in this
species: “(1) the tendency for these clams to develop
gametes and spawn progressively earlier in the season
northward and southward from northern Massachusetts,
and (2) the bimodal nature of spawning south of Cape
Cod.” They further indicated that the bimodality may
not have resulted from two distinct reproductive cycles,
but rather from an interruption in the spawning during a
single cycle. Bimodality of spawning appears to occur
with Mulinia lateralis also. Another possible reason for
the appearance of high numbers of Mulinia lateralis lar-
vae in late summer may be that some of the larvae which
had set during the earlier peak had become sexually
mature and spawned during the summer. I noted several
times during this study that under laboratory or natural
conditions little time was required for a new generation of
M. lateralis to develop gametes. Under natural conditions
I determined that these clams develop gametes within 6
weeks after setting. Some animals found in outdoor tanks
in late August, which had entered the tanks as free-
swimming larvae in early July and then metamorphosed,
had grown to sexual maturity. These specimens appeared
to be ripe and I was able to spawn them in the laboratory.
They may also have been capable of spawning under
natural conditions.
The possibility, however, that populations of Mulinia
lateralis from different areas belong to different physio-
logical races is not excluded. NELson (1928a,. 1928b)
was of the opinion that populations of American oysters,
Crassostrea virginica (GMELIN, 1791), which inhabit the
Atlantic Coast and the Gulf of Mexico were composed of
individuals of the same physiological race, or variety,
showing the same physiological requirements, such as the
minimum temperature at which they could begin spawn-
ing. Later studies by Coz (1934), LoosaANorF & ENGLE
(1942), LoosanorF & Tomm_rs (1948), SrauBER (1950),
and LoosanorF & NoMEjko (1951) suggested the existence
of physiologically different groups within the general popu-
lation of American oysters. LoosaNorF (1969) studied
the gametogenesis of different groups of American oysters
from Long Island Sound, New Jersey, Virginia, South
Carolina, and Florida by keeping them in Milford Har-
bor, Connecticut, for about 3 months and then subjecting
them to a long conditioning period at temperatures of
12°, 15°, or 18°C. The sharp differences in the rate of
development of their gonads caused him to conclude
that there are physiologically distinct populations of oys-
ters that require different temperature regimes for com-
pletion of gametogenesis and spawning.
Explanation of Plate 38
Section of Gonad Tissue of Female Mulinia lateralis
Figure 7: Active phase of oogenesis.
Figure 8: Fully mature. (XX 200)
Figure 9: Fully mature. - (X 500)
(X 500)
Figure 10: Partially spawned. (XX 200)
Figure 11: Spent, with few ova retained.
Figure 12: Inactive phase. (XX 200)
(X 200)
THE VELIcGER, Vol. 12, No. 3 [CALABRESE] Plate 38
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Vol. 12; No. 3
SUMMARY
1. Gametogenesis of Mulinia lateralis proceeded through-
out the year; the rate was low during October and Novem-
ber, immediately after spawning had occurred, and then
increased rapidly in December.
2. Of a total of 597 clams examined histologically, 302
(50.6% ) were females and 295 (49.4%) were males. No
hermaphrodites were found.
3. It was determined from histological sections that
spawning began in early July, when the water tempera-
ture increased to approximately 20° C, and continued into
September.
4, Larvae first appeared in plankton samples in early
July at water temperatures of 16° to 20° C and continued
to be in the plankton from early July to late October.
Peak abundance was in middle to late August, which co-
incided with the period when histological samples showed
greatest abundance of mature gametes.
5. It is probable that more than one generation of Mulinia
lateralis is produced in a single year.
ACKNOWLEDGMENTS
I thank the following members of the Milford laboratory
for their assistance: Dr. James E. Hanks and Mr. Harry
C. Davis for their many helpful suggestions throughout
this study and for their constructive criticism of this manu-
script; Mr. Manton L. Botsford for assistance in preparing
the photographs; and Miss Rita S. Riccio for her editorial
review.
LITERATURE CITED
BREUER, JOSEPH P.
1957. An ecological survey of Baffin and Alazan Bays, Texas.
Publ. Inst. Mar. Sci. Univ. Texas 4 (2): 134-155 (June ’57)
Cor, WesLEY RoswELL
1934. Alternation of sexuality in oysters. Amer. Natur.
68 (176): 236 - 251
Cronan, JoHN M., Jr.
1957. | Food and feeding habits of the scaups in Connecticut
waters. The Auk 74: 459 - 468
GiesE, ARTHUR CHARLES
1959. Comparative physiology: annual reproductive cycles of
marine invertebrates. In: Ann. Rev. Physiol. 21: 547 - 576.
V. E. Hall, ed. Ann. Rev., Inc., Palo Alto, Calif.
Hanxs, Rosert WILLIAM
1968. Benthic community formation in a “new” marine en-
vironment. Chesapeake Sci. 9 (3): 163-172 (Sept. 68)
THE VELIGER
Page 269
Humason, GretcHen L.
1962. Animal tissue techniques.
San Francisco. 468 pp.
Loosanorr, Victor LYon
1937. | Development of the primary gonad and sexual phases
in Venus mercenaria LINNAEUS. Biol. Bull. (Woods Hole)
72 (3): 389-405 (June 1937)
1969. Maturation of gonads of oysters, Crassostrea virginica,
of different geographical areas subjected to relatively low tem-
peratures. The Veliger 11 (3): 153 - 163; plts. 19 - 25
(1 January 1969)
Loosanorr, Victor Lyon « James B. ENGLE
1942. Accumulation and discharge of spawn by oysters living
at different depths. Biol. Bull. (Woods Hole) 82 (3):
413 - 422 (June 1942)
LoosanorF, Victor Lyon & CHartes A. NomEj]Ko
1951. Existence of physiologically-different races of oysters,
Crassostrea virginica. Biol. Bull. (Woods Hole) 101 (2):
151-156; 2 text figs.; 1 table (October 1951)
LoosanorF, Victor L. « FRANCES D. ToMMERS
1948. Effect of suspended silt and other substances on rate of
feeding of oysters. Science 107: 69 - 70 (16 Jan. ’48)
Loosanorr, Victor Lyon, Harry Caru Davis & PAuL E. CHANLEY
1966. Dimensions and shapes of larvae of some marine bivalve
mollusks. Malacologia 4 (2): 351 - 435; 61 text figs.
Netson, THURLOW CHRISTIAN
1928a. On the distribution of critical temperatures for spawn-
ing and for ciliary activity in bivalve molluscs. Science
67: 220 - 221
1928b. Relation of spawning of the oyster to temperature.
Ecology 9: 145 - 154
Ropes, Joun W.
1966. | Hermaphroditism in the surf clam, Spisula solidissima.
Ann. Reprt. Amer. Malacol. Union, p. 26
1968a. Reproductive cycle of the surf clam, Spisula solidissima,
in offshore New Jersey. Biol. Bull. (Woods Hole) 135 (2) :
349 - 365 (October 1968)
1968b. Hermaphroditism in the surf clam, Spisula solidissima.
Proc. Nat. Shellfish. Assoc. 58: 63 - 65 (June 1968)
Ropes, JoHN W. & ALDEN P. STICKNEY
1965. Reproductive cycle of Mya arenaria in New England.
Biol. Bull. (Woods Hole) 128 (2): 315 - 327 (April 1965)
SHaw, WILLIAM N.
1965. Seasonal setting patterns of five species of bivalves in
Tred Avon River, Maryland. Chesapeake Sci. 6 (1) : 33-37
(March 1965)
W.H. Freeman and Co.,
StauBer, LESLrE A.
1950. The problem of physiological species with special ref-
erence to oysters and oyster drills. | Ecology 31(1): 109-118
2 text figs.; 1 table (January 1950)
SuLLivaN, CHarLotTEe M.
1948. _ Bivalve larvae of Malpeque Bay, PE. I.
Res. Brd. Canada 77: 1 - 36
VERRILL, Appison E.
1873. | Report upon the invertebrate animals of Vineyard
Sound and the adjacent waters, with an account of the
physical characteristics of the region. Reprt. U.S. Fish
Comm. on the condition of the sea fisheries of the South Coast
of New England in 1871 and 1872: 295 - 747
Bull. Fish.
Page 270
THE VELIGER
Vol. 12; No. 3
Three New Species of Muricacean Gastropods
from the Eastern Pacific
WILLIAM K. EMERSON
Department of Living Invertebrates, American Museum of Natural History
Seventy-ninth Street and Central Park West, New York, New York 10024
AND
ANTHONY D’ATTILIO
Natural History Museum, San Diego, California 92112
(Plates 39, 40; 4 Text figures)
AS A RESULT OF OUR LONG-TERM investigation of the west .
American Muricacea, three new species, two referable to
the Muricidae and one to the Coralliophilidae, recently
came to our attention. We are indebted to Mr. and Mrs.
André DeRoy of Santa Cruz Island, Galapagos Islands
and Dr. James H. McLean of the Los Angeles County
Museum of Natural History for making this material
available for study.
Dr. George E. Radwin of the Natural History Museum
of San Diego, California and Professor Masao Azuma of
Nishinomiya, Japan kindly provided data on the radulae.
MoriciDAE
Murexiella radwini EMERSON & D’ATTILIO, spec. nov.
(Plate 39, Figures 1, 2; Text figure 2)
Description: The holotypic shell is 33.5 mm in length,
with 7 convex postnuclear whorls; 24 unsculptured nuc-
lear whorls are preserved on the smallest of the two im-
mature paratypes. The surface of the shell is richly imbri-
cated, especially on the projecting spines; and it is colored
a warm fleshy violet. The aperture is moderately small,
ovate; the columellar edge is slightly erect below, and the
outer lip is strongly crenulated by the exterior spiral
sculpture. The axial sculpture consists of 5 varices which
are obliquely descending to the right. Six spiral cords are
present on the body whorl. The upper 3 are recurved; the
lower 3 extend outwardly but are not recurved. There are
2 cords on the spire, which are spinose at the varices. The
major spiral cords are composed of 1 major ridge and 2 to
3 minor continuous ridges on each side; their surfaces are
overlaid by a continuous pattern of numerous tile-like
flattened lamellae; at the outer edges of the ridges the
lamellae develop into strong laterally projecting scales.
In the area between the last two varices the spiral sculp-
ture is much diminished. On the lower varices approaching
the spiny extensions of the spiral cords, there are abrupt
deep channels which strongly separate the major varical
spines. On their apertural side the spines are open; be-
tween the spines, except for their distal projections, there
are numerous, undulating, erect, blade-like lamellae. ‘The
narrowly opened siphonal canal is obliquely oriented to
the left, is flattened for most of its length, and recurved
only distally. The canal is ornamented on the right by 2
long spines and on the left by the ends of the previous
siphonal canals; on the remaining area of the canal,
there are present only growth striae. p
The aperture within is colored lightly but richly violet.
Exteriorly the shell is colored a fleshy violet except for the
varices which are a pinkish tan.
No periostracum is apparent. The operculum is muri-
coid, with a basal nucleus. The radular dentition of the
new species is illustrated (Text figure 2), and we are also
illustrating (Text figure 1) the radular dentition of Mu-
Vol. 12; No. 3
Figure 1
Murexiella hidalgoi (CrossE, 1869)
Central tooth and a lateral tooth, “Blake” Station 272, off Barbados
(U.S. N. M. No. 87081) ; greatly enlarged
rexiella hidalgoi (Crosse, 1869; U.S. N. M. no. 87081),
the type of the genus Murexiella CLENCH & FARFANTE,
1945. The radular slides of both species were kindly pre-
pared by Dr. George E. Radwin, Curator, San Diego
Natural History Museum.
Types: The holotype (A. M.N.H.no. 155903), which
measures 33.5 mm in length, was dredged by André and
Jacqueline DeRoy in 100m at Tagus Cove, Isabella Is-
land, Galapagos Islands, on January 29, 1968 (type local-
Figure 2
Murexiella radwint EMERSON & D’ATTILIO, spec. nov.
Central tooth and a lateral tooth from paratype
(S. D. N. H. S. No. 51335); greatly enlarged
ity). In addition, the following paratypic material was
studied: One immature specimen, height 16.8 mm (S. D.
N. H.S. no. 51335), from which the radula was extract-
ed; and a smaller immature specimen, height 13.5 mm
(A. D’Attilio collection). Both paratypes were dredged
from the same locality and on the same date as the holo-
type.
Remarks: This distinctive species is as yet known only
from the material at hand. It is related most closely to
Murexiella hidalgoi: (Crosse, 1869), of the western At-
THE VELIGER
Page 271
lantic. Crosse’s species has a narrower shell with longer
spines. Murexiella diomedaea (Dax, 1908), from the
eastern Pacific, has a slender, more spinose shell. Large
specimens of Murexiella humilis (Broperip, 1833), es-
pecially those found in the Bay of Panama, have a more
rebust, heavier shell and differ also in the short, blunt,
more recurved spines, and in other sculptural details.
“Murex” galapaganus EMERSON & J)’ATTILIO, spec. nov.
(Plate 39, Figures 3 to 6; Text figures 3, 4)
Description: Shell is dull white, of moderate size (at-
taining 46+ mm in length). Whorls are 7, the early
ones somewhat obscure through erosion, with 6 varices
ornamented with long recurved spines. The whorls are
subangulate, and, on the extended spire, there are 2
eee
A ravi b
Figure 3
Figure 4
Figures 3 and 4
“Murex” galapaganus EMERSON & D’ATTILIO, spec. nov.
Figure 3: Central tooth and lateral tooth from a paratype (A.
D’Attilio coll.) ; X 300 (drawing courtesy of Prof. Azuma)
Figure 4: Operculum, outer and inner side; greatly enlarged
Page 272
spiral cords, one on the shoulder and one above the im-
pressed suture. On the body whorl there is a low rounded
spiral cord on the shoulder, 2 low spiral cords appear
below, and one additional, much weaker cord is just above
the canal. The spines occur as extensions of the spiral
cords at each axial varix. The axial varices extend over
the shoulder diagonally as rounded, cord-like structures
and are depressed midway, rising at their junction with
the next whorl. The aperture is ovate, of moderate size
and polished white; the edge of the apertural lip is ele-
vated from the labial varix by a trough; the parietal lip
is not free in the holotype due to immaturity. The largest,
paratypic specimen (Plate 39, Figures 5, 6) has the pari-
etal lip mostly erect and has 8 poorly developed apertural
denticles on the outer lip.
One immature paratype (A. M.N.H.no. 155907)
possesses 24 smooth, nuclear whorls.
Although not preserved on the holotype, possibly due
to erosion, a thin outer layer of chalky matter occurs on
the paratypes. This chalky material, which is microscop-
ically striate, overlies the harder calcite surface.
The operculum has a terminal nucleus, situated sub-
laterally (Text figure 4). The radula is illustrated in Text
figure 3.
Types: Holotype (A. M.N.H.no. 155906), dredged in
150m, South Academy Bay, Santa Cruz Island, Gala-
pagos Islands, June 11, 1968, collected by the DeRoys.
Two paratypes (one in the DeRoy collection, length 43
mm; one in the D’Attilio collection, length 46mm) from
the same depth and locality, collected by the DeRoys,
June 10, 1968 (type locality).
Other paratypic material: One immature specimen, A.
M.N. H. no. 155907, dredged in 200 m, North Santa Cruz
Island, collected by the DeRoys, December 9, 1968, 11.5
mm in length. One immature specimen, Los Angeles
County Museum, Allan Hancock no. 816-38, north of
Hood Island, Galapagos Islands, 15 mm in length.
Remarks: On the basis of our present knowledge, we can
not assign with certainty this distinctive new species to an
existing genus. In shell morphology, it is similar to the
THE VELIGER
Vol. 12; No. 3
type species of Bathymurex CLENCH & TURNER, 1945, and
Paziella JoussEAUME, 1880, the radular characters of
which are not known. On the other hand, the radular
dentition of the type species of Takia Kuropa, 1953 (p.
190; fig. 10 on p. 180) is morphologically close to that of
the new species, but these taxa differ greatly in shell
morphology, especially in the lack of spines on the shells
of Takia. Both, however, have a chalky outer surface. Two
species described from Japanese waters, ““Boreotrophon”
gorgon Dau, 1913 and “B.” echinus Dati, 1918 have
similar opercula and radulae (Azuma, in litt.), but the
shells lack well-produced spines and an outer chalky layer.
Dall’s taxa were placed in Trophonopsis Bucguoy, DauT-
ZENBERG & DoLLFus, 1882, by Kuropa & HaBe (1952),
and were assigned to Bathymurex as a subgenus of Tro-
phonopsis by AzumMA (1960). These species were subse-
quently assigned to Paziella, with Bathymurex as a syno-
nym, by Voxes (1964). Thus, the new species could be
placed in the Trophoninae, Muricinae, or with the aspella-
like forms near Takia, depending on the generic assign-
ment one would choose to select for it. Although the Gala-
pagan specimens have a “trophonoid” appearance, we
hesitate to place the new species generically until we have
a better understanding of the taxonomic placement of
these generic taxa. If the new species were to be referred
to an existing genus on the basis of shell characters,
Paziella would appear to be the most promising assign-
ment.
CoRALLIOPHILIDAE
Latiaxis (Babelomurex) santacruzensis
EMERSON & D’ATTILIO, spec. nov.
(Plate 40, Figures 1 to 4)
Description: Shell of medium size, light in structure;
pale, ochre colored on its dorsum, aperture rosy pink:
Nucleus in the holotype is lacking and the first and per-
haps second whorl, in addition, are eroded. There follow
5 strongly carinated whorls each bearing on the carina
Explanation of Plate 39
Figures 1 and 2: Murexiella radwint EMERSON & D’ATTILIO,
new species, holotype; X 2
Figures 3 to 6: “Murex” galapaganus EMERSON & D’ATTILIO
§ g £4 >
new species
Figures 3, 4: Holotype (A. M. N. H. No. 155906); 2
Figures 5,6: Paratype (D’Attilio collection) from type locality; X 2
THE VELIGER, Vol. 12, No. 3 [Emerson & D’Arrtizio] Plate 39
Figure 1
i
t
H
i
Figure 4 Figure 5 Figure 6
Vol. 12; No. 3
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Page 273
extended, upturned, narrowly triangular spines; there are
10 of these long spines, some of which have been broken
off the shoulder carina; the number of spines per whorl
increases on each succeeding whorl. The whorls are deeply
parted at the suture, added to which the spiny peripheral
keel sharply emphasizes the angulated character of the
spire. Spiral sculpture is further evident in the presence of
weak threads on the spire which are axially overlaid by
poorly developed scabrous lamellae. This spiral sculpture
is present also on the convex surface of the body whorl
above the shoulder to about midway on the slightly
swollen body whorl. At the periphery of the body whorl
(but in an apertural view, at the junction of outer lip
and aperture), there are about 10 stronger spiral cords
intercalated with weaker cords extending anteriorly. Scab-
rous ornamentation overlies the spiral sculpture. Except
for 2 or 3 strong axial plicae nearer the outer lip, there
are only weak axial undulations on the body whorl.
The aperture is comparatively large, ovate, somewhat
angled at the juncture with the body; the lip is thin,
crenulated by reason of the exterior sculpture; a pro-
nounced groove is formed by the hollow edge of the spiral
keel. A small portion of the inner lip is adherent above but
erect below. The canal is strongly recurved with a coarsely
sculptured siphonal fasciole formed by the presence of 5
previous, preserved canals; a narrow but deep umbilical
chink is present.
Types: The holotype (A. M.N.H.no. 155901) is an
adult, 33.8 mm in length, greatest width 33.5 mm includ-
ing spines, dredged alive by the DeRoys, in 150 m, south
of Academy Bay, Santa Cruz Island, Galapagos Islands,
May 29, 1968 (type locality).
Paratypes: Anthony D’Attilio collection, one adult speci-
men; one immature specimen, 7.8 mm in length, A. M. N.
H. no. 155902; one immature specimen; all dredged with
the holotype. Jacqueline DeRoy collection, 1 mature spe-
cimen, 28 mm in length, dredged with the holotype (Plate
40, Figures 3, 4). One immature specimen, Los Angeles
County Museum, Allan Hancock no. 810-38, off Barring-
ton Island, Galapagos Islands, in 134 m.
Remarks: An unfigured paratype, no. 155902, A. M.N.
H. collection, retains the nucleus and protoconch in a
well-preserved condition. In this specimen there are 14
smooth nepionic whorls followed by 34 post-nepionic
whorls which have numerous erect lamellae. Each whorl
has a strong spiral shoulder ridge, above which the whorl
is slightly and convexly sloping toward the suture; it is
sculptured with the above mentioned blade-like lamel-
lae; below the shoulder ridge there is a second ridge close
to the suture; the area between the two ridges is convex;
the lamellae in this area continue from above but are
diagonally placed with respect to those above the shoulder.
There are present in the eastern Pacific a few represent-
atives of this genus. Only one of these, Latiaxis oldroydi
(Oxproyp, 1929) is of comparable size. Latiaxis oldroydi
is known to occur off southern California. The shell is
more robust, is white throughout, with coarser sculpture;
its spire is higher and with a narrower peripheral spiny
keel. Of more common occurrence in the Galapagos Is-
lands is Latiaxis hinds (CarPENTER, 1857) (= Tro-
phon muricatus Hinps, 1843). This species is collected in
shallow water and it has a shell of smaller size. Compari-
son may be made with one other species, Latiaxis dalli
EMERSON & D’Arttiio, 1963, from the western Atlantic.
Latiaxis dalli is nearest to L. santacruzensis, spec. nov., in
general form and size, differing in being entirely white
and in having well-developed, evenly dispersed spiral
cords over the entire shell.
LITERATURE CITED
Azuma, Masao
1960. A catalogue of the shell-bearing Mollusca of Okinoshimna,
Kashiwajima and the adjacent area (Tosa province) Shikoku,
Osaka, Japan. pp. 1-102; 1-17; plts.1-5 (20 March)
BropDERIP, WILLIAM JOHN
1833. | Characters of new species of Mollusca and Conchifera
collected by Mr. Cuming. Proc. Zool. Soc. London for
1832 (prt. 2): 173-179 (14 January 1833)
Bucguoy, E., PH. DauTzENBERG & G. F DotiFus
1882. | Les mollusques du Roussillon. Paris, vol. 1, fasc. 1:
1-40; plts. 1-6
CarPENTER, PHILIP PEARSALL
1857. | Report on the present state of our knowledge with
regard to the Mollusca of the west coast of North America.
Rep. Brit. Assoc. Adv. Sci. for 1856: 159 - 368; plts. 6-9
CiencH, WituiaM James « I. Pérez FaRFANTE
1945. The genus Murex in the western Atlantic.
Johnsonia 1 (17): 1-58; plts. 1-29 (29 May 1945)
Crosse, H.
1869. Diagnoses molluscorum novorum.
Paris, ser. 3, 9 (17) : 408 - 410
Dati, WILLIAM HEALEY
1908. Reports on the dredging operations off the west coast
of Central America to the Galapagos, to the west coast of
Mexico, and in the Gulf of California XIV. The
Mollusca and Brachiopoda. Bull. Mus. Comp. Zool., Har-
vard; 43 (6): 205 - 487; plts. 1-22 (October 1908)
1913. | Diagnoses of new shells from the Pacific Ocean. Proc.
U.S. Nat. Mus. 45 (2002) : 587 - 597 (11 June 1913)
1918. Notes on Chrysodomus and other mollusks from the
North Pacific Ocean. Proc. U.S. Nat. Mus. 54 (2234):
207 - 234 (5 April 1918)
Journ. Conchy]l.
Page 274
THE VELIGER
Vol. 12; No. 3
Emerson, WILLIAM KeitH & ANTHONY D’ATTILIO
1963. A new species of Latiaxis from the western Atlantic.
Amer. Mus. Novitates no. 2149: 1 - 9; 6 text figs.
(25 July 1963)
JoussEAuME, FELIx PIERRE
1880. Division méthodique de la famille des purpuridés.
Le Naturaliste 2 (42) : 335 - 336
Kuropa, TOKUBEI
1953. On the Japanese species of “Trophon.”
(4): 186 - 202; text figs. 1-8
Venus 17
Kuropa, ToKuBEI & TADASHIGE HABE
1952. Checklist and bibliography of the Recent marine Mol-
lusca of. Japan. 210 pp.; 2 maps. Tokyo, Japan (4 April 1952)
Otproyp, IpaA SHEPARD
1929. Description of a new Coralliophila.
42 (3): 98, 99; plt. 5
Vokes, Emity Hoskins
1964. Supraspecific groups in the subfamilies Muricinae and
Tritonaliinae (Gastropoda: Muricidae). Malacologia 2 (1):
1-41; plts. 1-3 (September 1964)
The Nautilus
(January 1929)
Explanation of Plate 40
Latiaxis (Babelomurex) santacruzensis EMERSON & D’ATTILIO,
new species
Figures 1,2: Holotype (A. M. N. H. No. 155901); X 2
Figures 3,4: Paratype (J. DeRoy collection) , from type locality; X 2
THE VELIGER, Vol. 12, No. 3 [EMERSON & D’ArrTiLio] Plate 40
Figure 1 Figure 2
Figure 3
Vol. 12; No. 3
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Page 275
A New Species of Helminthoglypta from the Mojave Desert
WALTER B. MILLER
Department of Biological Sciences, University of Arizona, Tucson, Arizona 85721
(Plate 41; 2 Text figures)
In 1930, S.S. BERRY DESCRIBED a new species of desert
snail from the El Paso Mountains of the northern Mojave
desert of California. With only shell characters for dia-
gnosis, he named it Micrarionta (Eremarionta) microme-
talleus S.S. Berry, 1930. Subsequently he dissected adult
specimens and determined that the anatomical characters
were those of Sonorella. The shell characters, however,
were different from those of other known Sonorella, and
in 1943 he established a new subgenus Mohavelix to ac-
commodate species of Sonorella with small, thin, subdis-
coid, and widely umbilicated shells. Accordingly, M. (E.)
micrometalleus became Sonorella (Mohavelix) microme-
talleus (BERRY), the type and only species of Mohavelix.
Since 1964 I have been intensively studying Sonorella
and in 1967 I submitted a revision of the genus to the
University of Arizona as my doctoral dissertation. I be-
came increasingly convinced that Sonorella (Mohavelix)
micrometalleus did not share a close phylogeny with other
Sonorella, i. e., they did not evolve from the same immedi-
ate pre-Sonorella ancestor, and eventually proposed (Mit-
LER, 1968) that Mohavelix should be raised to generic
rank. The phylogeny of Mohavelix has remained obscure,
however, with the most credible hypothesis being its deri-
vation from some ancestral Eremarionta, possibly close to
Eremarionta aquaealbae Berry, 1922.
While attempting to obtain live specimens of Mohavelix
micrometalleus from the arid southern slopes of the El
Paso Mountains (the type locality is in the southern part
of Last Chance Canyon) in early January 1969, I decided
to search for more suitable rockslides on the more humid
northern slopes of the range. I was rewarded by finding a
gigantic north-facing slide which yielded 95 dead shells
and 10 live specimens, of which 3 were adult. They looked
in all respects like M. micrometalleus. Jubilant over this
new find in a prolific locality, I was totally astonished to
find that the anatomy revealed these snails to be a new
species of Helminthoglypta, described below. The discov-
ery of this population of Helminthoglypta in the El Paso
Mountains, with shell and certain anatomical characters
remarkably similar to those of M. micrometalleus, has
now afforded a more credible hypothesis for the derivation
of this species. This hypothesis is also discussed below.
Helminthoglypta micrometalleoides W. B. Mitirr,
spec. nov.
(Plate 41, Figures 1 and 2; Text figure 1)
Description of Holotype: Shell very small for the genus,
depressed, discoid, thin, light-brown, with a darker brown
spiral band on the well-rounded shoulder; widely umbili-
cate, the umbilicus contained about 6 times in the diame-
ter of the shell. Embryonic shell of about 13 whorls, with
faint, microscopic wrinkles. Post-embryonic whorls with
minute granular wrinkles and papillae. Body whorl with
spirally-descending, long, hyphen-like papillae occasion-
ally confluent to form long threads, superimposed over
the radial ridges, persisting into the umbilicus; periostra-
cum thin, silky-lustrous. The last whorl descends slightly
to the scarcely expanded, thin peristome; aperture oblique,
relatively large.
Shell Measurements: Height 5.6 mm, maximum diame-
ter 10.9 mm, umbilicus 1.7 mm. Number of whorls 44.
The animal: The animal, when extended, has a dark-grey
to black body wall, with scattered white glandular papil-
lae. The mantle collar shows a thick mat of white mucus
glands, giving the collar an overall whitish appearance;
the area around the pneumostome is chalk-white. The
mantle above the lung is pigmented with small, black,
closely-spaced spots.
The Genitalia: The genitalia (Text figure 1) are typical
for the genus. The penis is strongly swollen in the middle
where the internal tube, detached from the external wall,
becomes thickened and forms a papilla which can be
likened to a very short verge. Anteriorly, it consists of a
hollow, eversible sac. Distally, it merges into the epiphal-
Page 276
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Vol. 12; No. 3
lus where the internal tube becomes completely adnate
to the external wall. The epiphallus is of uniform diame-
ter and is equipped with a moderately long epiphallic
caecum at its distal end. The penial retractor muscle
originates on the floor of the lung and is inserted on the
epiphallus near the penial end. The vagina is short, equal
hd
ag)
5.0 mm
in length to the saccular proximal end of the penis. The
free oviduct is longer than the vagina. The long sperma-
thecal duct gives rise to a diverticulum about halfway
along its length; the diverticulum is about equal in length
to that part of the spermathecal duct posterior to their
junction. The globular spermatheca is bound by connec-
tive tissue to the posterior end of the uterus. The muscu-
lar dart sac is small, globular, and situated at the summit
of a longer atrial sac; it contains a short, conical dart. The
two side-by-side mucus bulbs join proximally into a single
duct which enters the atrial sac at its junction with the
dart sac; distally, each mucus bulb connects with a thin,
glandular, U-shaped duct which passes into a broad, thin
membrane enveloping the entire dart apparatus and much
of the lower genitalia. Dimensions, in millimeters, of dis-
tinctive structures follow:
Penis 5.0 mm
Epiphallus 7.0 mm
Epiphallic caecum 8.0 mm
Penial retractor 3.5mm
Spermathecal duct 14.5 mm
Spermathecal diverticulum 8.5 mm
Vagina 2.0 mm
Free oviduct 3.0 mm
Type Locality: El Paso Mountains, Kern County, Califor-
nia, in north-facing, high rockslide of small rocks, between
crag outcroppings on south side of Iron Canyon Road,
at a point 3 miles up the canyon from the junction of the
road with the Garlock-Goler highway (W.B. Miller, 5
January 1969).
Disposition of Specimens:
Holotype: deposited in the California Academy of
Sciences, Geology Type Collection, no. 13169.
Figure 1
(<— adjacent column)
Helminthoglypta micrometalleoides W. B. MiLuEr, spec. nov.
Lower genitalia of holotype; drawing made from projection of
stained whole mount. Scale in millimeters >
ag albumen gland ats atrial sac da dart
dp distal part of penis ds_ dart sac
ec epiphallic caecum ep epiphallus fo free oviduct
hd hermaphroditic duct
itp internal tube of penis mb mucus bulb
pp proximal part of penis prm penial retractor muscle
prs prostate sp spermatheca spd spermathecal duct
spdv spermathecal diverticulum ut uterus
va vagina vd vas deferens
go genital orifice
Tue VELIGER, Vol. 12, No. 3
[MiLiER] Plate 41
Figure 1 Figure 2
esl ne ll
Figure 3 Figure 4
Figures 1, 2: Helminthoglypta micrometalleoides W. B. Mier,
spec. nov. Holotype. California Academy of Sciences, Geology Type
collection no. 13169. El Paso Mountains, Kern County, California.
Scale in millimeters
Figures 3, 4: Mohavelix micrometalleus (BERRY, 1930). El Paso
Mountains, Kern County, California. Scale in millimeters
Vol. 12; No. 3
Paratypes: in the Invertebrate Museum, Department
of Biological Sciences, University of Arizona, and in the
private collection of the author.
Remarks: Helminthoglypta micrometalleoides is the
smallest species of Helminthoglypta described to date.
Examination of approximately 100 paratypes (not all
adults) reveals a remarkable constancy of characters. The
maximum diameter of the shell varies only from 9.3 mm
in the smallest adult to 11.3 mm in the largest. Shell sculp-
ture depends on the age of the shell; older shells have
worn embryonic whorls and papillae. The umbilical di-
ameter varies from as large as to slightly larger than that
of the holotype. The genitalia from three dissected speci-
mens do not appear to provide diagnostic characters to
distinguish this species from other desert helminthoglypts.
The peculiarly shaped penis is a characteristic of all
known and dissected desert helminthoglypts, as reported
by W. O. Gregg (in litt.).
Helminthoglypta micrometalleoides appears to be most
closely related to H. fishert (BartscuH, 1904) of the Pan-
amint Mountains and to H. greggi WILLeTT, 1931, of
Soledad Mountain. It can be readily distinguished from
these other species by the much smaller shell diameter.
The El Paso Mountains are situated geographically be-
tween the above localities. In field diagnosis, H. micro-
metalleoides can be easily mistaken for Mohavelix micro-
metalleus. The size, shape, color, texture, and general ap-
pearance of the shell are the same for both species (Plate
41, Figures 3 and 4). Microscopic examination of fresh
shells reveals that M/. micrometalleus has an embryonic
sculpture of thickly-set, spirally arranged, hyphen-like
papillae similar to Eremarionta. The post-embryonic
sculpture is more papillose and radially wrinkled than
that of H. micrometalleoides and the periostracum is less
glossy. The adult genitalia provide the major diagnostic
difference between Helminthoglypta and Mohavelix. Mo-
havelix (Text figure 2) has simplified genitalia similar to
Sonorella. The atrial sac, dart sac mucus glands, and
spermathecal diverticulum are all missing, while the epi-
phallic caecum is reduced to a vestige; the penis has a
short verge at its distal end. It is most interesting to note,
however, that the remaining structures, such as penis,
epiphallus, spermathecal duct, vagina, free oviduct, and
uterus have the same relative dimensions as in H. micro-
metalleoides.
Concerning the phylogeny of Helminthoglypta micro-
metalleordes, it appears most likely that this species evolved
from a common ancestor of the desert helminthoglypts.
It is debatable that all described species of desert hel-
minthoglypts are good biological species, with fully estab-
lished reproductive isolation; hybridizing experiments be-
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Page 277
hd
ag
2
4
: |
=
2
Be spd i
prs 42: ut Hf
= V
it
Gr
ae
fo
ve
NS
bf er
cc,
= pe
go
5.0 mm
Figure 2
Mohavelix micrometalleus (BERRY, 1930)
Lower genitalia; drawing made from projection of stained whole
mount. Scale in millimeters
ag albumen gland ec epiphallic caecum
ep epiphallus fo free oviduct go genital orifice
hd hermaphroditic duct pe penis
prm penial retractor muscle prs prostate
sp spermatheca spd spermathecal duct ut uterus
va vagina vd vas deferens ve verge
Page 278
tween H. micrometalleoides, H. greggi, and H. fishert
would be desirable. The large difference in size of H.
micrometalleoides from all other known species of Hel-
minthoglypta would suggest reproductive isolation from
sheer mechanical incompatibility, even if other physio-
logical, genetic, or chromosomal barriers did not exist.
What appears to be most interesting and significant,
however, is the possible evolutionary relationship of
Helminthoglypta micrometalleoides to Mohavelix micro-
metalleus. The nearest relative of M. micrometalleus is not
known, although, as stated earlier, a descent from an
ancestral Eremarionta would be plausible; a possible de-
scent from an ancestral Sonorella is too difficult to support.
It is now suggested that M. micrometalleus evolved direct-
ly from an isolated population of H. micrometalleoides.
The close similarities of shell characteristics of H. micro-
metalleoides and M. micrometalleus, as well as the simi-
larities in relative size and shape of the significant ana-
tomical structures common to both species suggest a close
relationship. The evolution of the Helminthoglyptidae is
replete with instances of secondary simplification of ana-
tomical structures, as in the case of Sonorelix, Sonorella,
Tryonigens, Micrarionta, Eremarionta, as well as Mohav-
elix. The mechanism of this simplification has not been
determined. Populations of different genera and species
are often in similar ecological niches, sometimes sympatric,
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Vol. 12; No. 3
and the adaptive advantages of a simplified reproductive
system are not apparent. Genetic drift is the most likely
mechanism in desert populations where marginal isolates
are periodically and frequently subjected to prolonged
drought to the point where individual numbers become
critically low. Chromosomal breakage, inversion, reduc-
tion, or translocation could account for large losses of
structures, which, fortunately, are not fatal.
It is hoped that studies of the chromosomes of Hel-
minthoglypta micrometalleoides and Mohavelix micro-
metalleus can be undertaken in the near future. Also,
gross comparison of proteins by chromatography or elect-
rophoresis might yield data on the extent of relationship
between the two species.
The specific name is chosen to indicate the close resem-
blance between the two species.
LITERATURE CITED
BErry, SAMUEL STILLMAN
1930. New helicoid snails from the Mohave Desert. Ann.
Mag. Nat. Hist. 10 (6): 187 - 193
1943. | On the generic relationships of certain Californian xero-
phile snails. Trans. San Diego Soc. Nat. Hist. 10 (1): 1 - 24
MILLER, WALTER BERNARD
1968. New Sonorella from Arizona.
30 - 63
The Nautilus 82 (2) :
Vol. 12; No. 3
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Page 279
Two New Species of the Genus Caldukia Burn & MILLER, 1969
(Mollusca : Gastropoda : Opisthobranchia )
from New Zealand Waters
MICHAEL C. MILLER
Department of Zoology, University of Auckland, Private Bag, Auckland, New Zealand
(21 Text figures)
INTRODUCTION
THE GENUS Caldukia BURN & MILLER, 1969 was estab-
lished to embrace 3 species of janolid nudibranch, one
Australian and two from New Zealand, which could not
be placed in any of the known genera. Caldukia is distin-
guished from the other genera of the family JANOLIDAE
(Zephyrina QuaTREFAGES, 1844; Janolus Bercu, 1884;
Antiopella Hoye, 1902; a new genus (MILLER, in press)
by the shape of the rhinophoral club (short and stout witha
small series of primary lamellae, each with regularly dis-
posed rows of secondaries), the lack of an inter-rhino-
phoral crest, a radula of the formula 6-1-6 with the
lateral teeth strongly cuspidate, and very strong unequal
jaws with a few large horny teeth. A fairly complete
comparison of the genera has already been made (Mar-
cus, 1955, 1958; BurN & MILter, 1969; MILter, in press).
The Australian species, which is the type of the genus, was
first described a little over a decade ago and at that time
was provisionally assigned to the genus Proctonotus ALDER
& Hancock, 1844 (Burn, 1958) ; although the two New
Zealand species were first discovered in 1961, original
descriptions of them are only now being presented.
ACKNOWLEDGMENTS
I wish to thank Professor J. E. Morton for advice and
encouragement, Mr Robert Burn for discussing arminid
systematics and nomenclature and for letting me examine
his colour photograph of a living specimen of Caldukia
affinis, Dr E. J. Batham for permitting me to use the
Portobello Marine Biological Station and the University
Grants Committee (New Zealand) for financing much of
my field work.
CLASSIFICATION
NUDIBRANCHIA
Arminacea
PACHYGNATHA
JANOLIDAE
DESCRIPTIONS
Caldukia albolineata MitteR, spec. nov.
Morphology: Length, extended, 11 mm (only one speci-
men collected). Body soft, lanceolate in outline (anterior
end blunt) ; low, back slightly arched with a large ellip-
tical swelling, the pericardium, in the centre, which is
confluent posteriorly with a prominent longitudinal ridge
(the dorsal sinus) which runs part way to the anus
(Figure 1). Rhinophores one-sixth to one-seventh of the
length of the body; peduncle very short; club stout with 6
sloping primary lamellae (lamellae of the two sides of
the rhinophore widely separate anteriorly, but approxi-
mate posteriorly), each with an upper and lower row of
small plates arranged alternately, and the rachis is pro-
duced apically as a cylindrical truncate process about
one-third of the height of the rhinophore (Figure 2).
Oral tentacles are short, blunt triangular lobes. Cerata
linear, apices pointed, non-caducous and very mobile
(Figure 3) ; the largest, when extended, roughly one-fifth
of the length of the body; inserted in 3 staggered longi-
tudinal rows, cerata smallest at the edge of the notum,
increase in size centripetally (Figure 4). Foot rounded
at the front; tail short, tapers to a fine point. Anus at the
tip of a large papilla situated in the mid-line near the
Page 280
Figure 1
Caldukia albolineata
Dorsal view of the living animal
posterior end of the notum. Reproductive apertures on the
right side, just below the edge of the notum at a level
midway between the rhinophores and the front of the peri-
cardium. Renal pore on the right side, just below the edge
of the notum and at a little distance from the rear end of
the pericardium (Figure 4).
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Vol. 12; No. 3
Colour: Body translucent, pale dull yellow; hermaphrodite
and female accessory glands show through as a central
opaque mass. Rhinophores pale dull yellow, apical process
covered with opaque white. Each ceras with a dark brown
diverticulum, centre of inner surface with a narrow lumpy
stripe of opaque white running from the base to the tip,
flanked by fairly large spots of the same pigment (Figure
3). Dorsal surface of tail with a wedge-shaped patch of
opaque white.
Alimentary System: Figure 5. Oral tube short and wide,
leads to the massive, ovate (when viewed from above),
Figure 2
Caldukia albolineata
Side view of a rhinophore
Figure 3
Caldukia albolineata
Cerata of the living animal: left — view of inner surface,
right — view of lateral surface
Vol. 12; No. 3
THE VELIGER
Page 281
Figure 4
Caldukia albolineata
View of the right side of the body showing the insertions
of the cerata and the apertures
an = anus
dorso-ventrally flattened buccal bulb. Salivary glands
huge, folliculose, lie closely applied to the sides of the
oesophagus and ventro-lateral walls of the stomach (in
the one specimen examined the condition of the salivary
glands was probably abnormal, the left gland extended
behind the stomach along the main posterior duct of the
digestive gland and ended at the origin of the second
left lateral duct — the right gland was very small and
barely reached the stomach). Oesophagus short, wide and
muscular, runs from the centre of the upper surface of the
buccal bulb to the fairly large banana-shaped muscular
stomach which lies to the left of the mid-line. There are
3 main stomachal ducts of the digestive gland; the right
and left anterior ducts are short and each opens into a
wide longitudinal duct which extends along the edge of
the anterior half of the notum on the left side and two-
thirds on the right. Short lateral branches, simple and
divided, arise along the sides of the longitudinal duct, a
few on the inner side and many on the outer. The poste-
rior duct, which shares a common opening into the stom-
ach with the left anterior duct, is long; it runs along the
left side of the gonad and then, just before the hind end
of the pericardium, starts to bend over to the mid-line of
the body, giving off as it does so, two lateral ducts to the
left side. On reaching the mid-line, the main duct bends
sharply, giving off as it does the first right lateral duct, to
run posteriorly above the intestine, sending branches to
the left side. It veers gradually to the left side of the
anus and then curves to the right around the posterior
end of the notum sending off short branches; all but the
most posterior of the lateral ducts branch. Diverticula
arise singly from the longitudinal ducts and side branches.
rp = renal pore
ra = reproductive apertures
Each diverticulum is simple and extends almost to the tip
of the ceras (Figure 6). The intestine arises on the left
side from the posterior end of the stomach; it bends im-
mediately through 180° and then describes a half circle
around the front of the pericardium as it passes to the
right side. On the right side the intestine continues to
bend and passes below the posterior end of the pericardi-
um to return to the left; it then bends again to run to
the anus.
Buccal Armature: Radula (Figure 7). Formula 6-1-6;
the one specimen examined had a radula of 28 (including
6 developing) rows. Central tooth smallish, roughly tra-
pezoidal; posterior face concave, upper edge notched to
form a broad semilunar median cusp with a small apical
denticle; the notched edge is serrulate and there is a
denticle on each of the corners. The lateral teeth are
peg-like; the inner 5 (2 and 3 are the largest) are fairly
similar in shape with a broad cuspidate crown, the middle
cusp being very large, and a somewhat sinuous and slightly
tapered base (short in 5) bent towards the margin of the
radula; the sixth (the outermost) is very small, almost
rectangular with a single (median) cusp. Jaws (Figure 8).
Strong, unequal, united dorsally by a thick band of chitin.
Right jaw long, fairly narrow and curved with 4 large
horny teeth at the anterior end. Left slightly shorter than
the right, but broader and angular, with 2 large teeth.
Nervous System: Although worked out in some detail it
is not described here: the most important features are the
long optic nerves and the fused cerebral and pleural
ganglia.
Page 282 THE VELIGER . Vol. 12; No. 3
2) LY
BORK
hort
oe
vd
=
Kon V,
San
BA
Figure 6
Caldukia albolineata
Preserved ceras (stained with borax carmine and cleared with
cedar-wood oil) showing diverticulum of digestive gland
Kidney: Figure 5. This organ lies dorsally between the
front end of the pericardium and the anus: it is thin
walled and consists of a long central sac and a number
of lateral diverticula, some reaching almost to the edge
of the notum; the anterior diverticula are long and
irregularly lobed, those to the posterior are short and
simple. The renopericardial duct (organ) is small and
Figure 5
(<— adjacent column)
Caldukia albolineata
Dorsal view of the general anatomy
an = anus bb = buccalbulb cns = central nervous system
he = heart in = intestine ki = kidney
Idg = left digestive gland ot = oraltube pc = pericardium
pe = penis rd = renopericardial duct
rdg = right digestive gland rp = renal pore
sg = salivary gland st = stomach
Wolw 12: No. 3 THE VELIGER Page 283
AS |
Figure 7
Caldukia albolineata
Radular teeth, half row
C = central (rhachidian) Li = innermost lateral
L6 = outermost lateral
links the pericardium with the first large right diverticu-
lum. The renal pore opens at the tip of the last large
right diverticulum.
Reproductive System: Figure 9. Hermaphrodite gland of
4 follicles, each joins separately, by a long ductule, to the
common hermaphroditic duct which runs forward to the
ampulla. The ampulla is ovoidal with a small chamber
(or pouch) on the right side into which opens the herm-
aphrodite duct and, by a short narrow duct, a small
vesicle (the fertilization chamber). The vas deferens and
oviduct arise separately, the former from the front end of
the ampulla, the latter from the front end of the side
chamber. The first third of the vas deferens is a narrow
duct, the remainder is a very wide glandular portion (the
prostate) ; the penis is conical and unarmed (Figure 10).
The oviduct runs to the right, constricts and then forks,
one of the branches leads to the albumen gland, the other
to the female atrium. The bursa copulatrix is ovoid and
is connected to the atrium by a long narrow duct, the
vagina.
Figure 8 Locality and Habitat: New Zealand, the South Island:
Caldukia albolineata Otago Harbour, Aquarium Point, one specimen on the
Jaws, view of inner surface under surface of a rock in the sublittoral fringe, 20
upper — right Jaw lower — left jaw January 1961.
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THE VELIGER
Vol. 12; No. 3
Figure 9
Caldukia albolineata
Reproductive system, unravelled
am = ampulla be = bursa copulatrix fa = female atrium
fe = fertilization chamber fgm = female gland mass
hd = hermaphrodite duct hg = hermaphrodite gland
ov = oviduct po = pouch pr = prostate
ps = penial sheath va = vagina vd = vas deferens
Type: Holotype — a microscope slide of the radula and
a colour photograph (35 mm Kodachrome transparency)
of the one living animal collected at Aquarium Point,
deposited in the Dominion Museum, Wellington (M.
PATA)
Caldukia rubiginosa Mi.tEr, spec. nov.
Morphology: Figure 11. Extended length up to 12 mm.
As the previous species except for the following details:
rhinophores with up to 7 lamellae, lowest very small (Fig-
ure 12) ; cerata fusiform (Figures 13, 14), in up to 4 rows.
Colour: Dorsal part of the body, rhinophores, distal por-
tion of cerata, opaque reddish brown; cerata transparent
(except at base), upper half yellow, lower half colourless,
latter divided by a band of opaque white (with blue iri-
descence) which is broad to the centre of the body and
narrow to the outside, diverticula apricot or pale brown;
upper region of the side of the foot speckled with reddish
brown, lower region, tail and sole transparent yellow or
apricot — hermaphrodite gland visible as opaque pink
bodies.
Vol. 12; No. 3
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Page 285
Figure 10
Caldukia albolineata
Penis, preserved (stained with borax carmine, cleared in
cedar-wood oil)
pe = penis ps = penial sheath pr = prostate
Alimentary System: As Caldukia albolineata spec. nov.
There is, however, a right post-anal branch of the digestive
gland.
Buccal Armature: Radula (Figure 15). A 9mm animal
had a radular formula of 23 (including 5 developing rows)
x 6:1-6. Teeth very similar to those of C. albolineata:
central with weak serrulations, central cusp without an
apical denticle, no denticles at the corners; laterals rather
uneven in outline, inner 4 teeth with a small flange de-
veloped on both the inner and outer sides of the central
cusp. Jaws (Figure 16): as in C. albolineata, but the right
jaw is slightly more angular.
Kidney: Large, filling the dorsal region of the body cavity
from the middle of the pericardium to the anus. More
branched than in C. albolineata and the branches are so
compacted as to give the organ an alveolate appearance.
Reproductive System: Figures 17, 18. Very similar in plan
to that of Caldukia albolineata. However, the ampulla is
oblong, the hermaphrodite duct does not enter by way of
a lateral swelling and there is no vesicle (fertilization
chamber) at the side; also, a short, narrow, common duct
leaves the front end of the ampulla and this bifurcates
into vas deferens and oviduct.
Locality and Habitat: New Zealand, the North Island:
Goat Island Bay, near Leigh; on the under surfaces of
rocks in the sublittoral fringe, feeding on the polyzoan
a
Figure 11
Caldukia rubiginosa
Dorsal view of a living animal
Beania magellanica (Busk) ; six specimens, one spawn
band, 28 August 1961; two specimens, several spawn
bands, 26 October 1961; two specimens, 26 February
1963.
Page 286 THE VELIGER ) Vol. 12; No. 3
Type: Holotype: a specimen collected at Goat Island Bay,
near Leigh, deposited in the Dominion Museum, Welling-
ton, New Zealand (M. 22178).
Food and Feeding Habits:' Caldukia rubiginosa feeds
on the polyzoan Beania magellanica (BusK) which it
tears apart with its large, strong dentate jaws. A single
zooid is attacked — this and adjacent zooids are first
smothered with mucus. When feeding, the mouth is opened
wide and the buccal bulb thrust forward so that the jaws
and radula are protruded (Figure 20). Jaw and radular
movements are synchronized with those of the buccal
bulb; when the latter is thrust forward the jaws and the
two halves of the radula open, when withdrawn, they
close. The polyzoan zocecium, which is usually grasped
Figure 12 about the middle, is fractured by the large horny teeth
Caldukia rubiginosa
Side view of a rhinophore ™ Mentioned briefly in Morton « MILLER (1968) on p. 412 but
the species is not named
Figure 13 Figure 14
Caldukia rubiginosa Caldukia rubiginosa
Cerata of a living animal; left drawing shows pigmentation and Preserved ceras (stained with borax carmine and cleared with
diverticulum of the digestive gland cedar-wood oil) showing diverticulum of the digestive gland
C Li
_eenee re Figure 15
Caldukia rubiginosa
Radular teeth, half row
C = central (rhachidian) Li = innermost lateral
L6 = outermost lateral
Vol. 12; No. 3 THE VELIGER
Imm
Figure 16
Caldukia rubiginosa
Jaws, view of inner surface
upper — right jaw lower — left jaw
of the jaws closing upon it and drawing it into the buccal
cavity, assisted by a sharp contraction of the head on the
firmly fixed foot. This is repeated many times until the
zooid, or part of it, is torn from the colony (Figure 21).
The polypide is completely macerated during the tearing
process. The radula acts as a conveyor belt and transports
the detached zooid, or fragment, to the oesophagus. The
whole action is very rhythmical, there being an active
phase of 10 - 25 seconds during which the animal makes
4 to 8 thrusts with its buccal mass, followed by a resting
phase of 60 - 70 seconds. The nudibranch may take up to
35 minutes to detach a single zooid. The zocecia of the
zooids ingested are not altered as they pass through the
Figure 17
Caldukia rubiginosa
Page 287
Reproductive system, unravelled
be = bursa copulatrix
ampulla
fgm = female gland mass
hg = hermaphrodite gland ov = oviduct
ps = penial sheath
pe
va = vagina
Figure 18
Caldukia rubiginosa
hd = hermaphrodite duct
pr = prostate
vd = vas deferens
Penis, preserved (stained with borax carmine, cleared in
pe = penis
cedar-wood oil)
ps = penial sheath
pr = prostate
Page 288 THE VELIGER Vol. 12; No. 3
a
Figure 19
Caldukia rubiginosa
Drawing of an animal feeding on the polyzoan
Beania magellanica (Busx)
Figure 20
Caldukia rubiginosa
Ventral view of the head of a living animal showing the buccal
bulb thrust through the dilated mouth and the jaws and two halves
of radula opened
gut; they are, however, compacted into a firm string
before being defaecated.
Figure 21
Caldukia rubiginosa
DISCUSSION Series of sketches showing an animal (head only shown) biting off
a single zooid of Beania magellanica (Busk).
é We ; Bottom — a ventral view of head with an avicularium (bitten off
Colour and colour pattern clearly distinguish the two with the zooid) protruding from the mouth. A, B, C — letters
species of Caldukia, described here, from each other and identifying the zooids
Vol. 12; No. 3
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Page 289
from the type species, C. affinis (Burn). Briefly stated:
C. rubiginosa has a reddish brown body and cerata with
colourless, yellow and opaque white (blue iridescence)
zones; C’. albolineata has a dull yellow body and a single
opaque white line down each ceras; and C. affinis has a
reddish fawn body marked mid-dorsally with yellow and
fawn cerata speckled with minute red spots. The colours
and patterns appear to be constant in C. rubiginosa and
C. affinis. There are only minor morphological and ana-
tomical differences between the three species. Most im-
portant of these is the shape of the radular teeth, par-
ticularly the central and the first and sixth laterals. In C.
albolineata the central is broad with a wide median cusp,
which, like the corners, is apiculate and serrulate. This
tooth in C. rubiginosa differs in being longer (when viewed
from above) and without the denticles on the cusp and the
corners (though these may have been worn off in the
animal examined). Caldukia affinis has a central tooth
with a narrow blunt cusp and no denticles or serrulations.
The first lateral tooth of C. affinis is almost rectangular,
but in the other two species it has a comma-shaped base
(like lateral teeth 2 to 5 of all 3 species); the median
cusps of the first to fourth lateral teeth of C’. rubiginosa
have small side flanges. The sixth tooth of C. affinis has
a fairly long, pointed and bent base, whereas in the other
2 species it is very short and blunt. The jaws differ only
slightly in outline. In the few specimens examined there
were minor variations in the branching of the digestive
gland, but these did not appear to be constant enough to
be used to separate the species. The reproductive system
is fundamentally similar in the 3 species; however, that
of C. albolineata differs from the systems of the other 2
species in the following 2 respects: (1) the male and
female ducts leave the ampulla separately, i.e., there is
no anterior common duct, and (2) the fertilization cham-
ber is a vesicle at the side of the ampulla.
SUMMARY
Two new species of the janolid nudibranch genus Caldukia
Burn & MILLER are described, viz. C. albolineata and C.
rubiginosa. The anatomy of C’.. albolineata is given in some
detail and the feeding habits of C’. rwbiginosa, which preys
on the polyzoan Beania magellanica (Busk), are briefly
described. The two new species and the type of the genus,
C. affinis (BURN) are compared; colour and pattern are
the principal distinguishing characteristics; small differ-
ences in the radular teeth and reproductive system are of
secondary importance.
LITERATURE CITED
Burn, RoBERT
1958. Further Victorian Opisthobranchia.
Soc. Australia 2: 20 - 36
Burn, Ropert & MicHAEL CHARLES MILLER
1969. A new genus, Caldukia, and an extended description
of the type species, Proctonotus ? affinis Burn, 1958 (Mollus-
ca: Gastropoda: Arminacea, Antiopellidae). Journ. malacol.
Soc. Australia 12: 23 - 31
Marcus, ERNST
1955. | Opisthobranchia from Brazil. Bol. Fac. Fil. Univ.
Sao Paulo, Zoologia 20: 89 - 200; plts. 1 - 30
1958. | On Western Atlantic opisthobranchiate gastropods.
American Mus. Novitates no. 1906: 1 - 82
Miter, MicHAEL CHARLES
in press, A new genus and species of the nudibranch family Ja-
nolidae (Mollusca; Gastropoda, Opisthobranchia) from New
Zealand waters. Journ. nat. hist. London
Morton, JoHN Epwarp & MicHAEL CHARLES MILLER
1968. The New Zealand sea shore. Collins, London and
Auckland; 638 pp.
Journ. malacol.
Page 290
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Vol. 12; No. 3
Occurrence of a Rare Squid, Chaunoteuthis mollis APPELLOF
(Family Onychoteuthidae) in the Indian Ocean
BY
K. N. NESIS
Institute of Oceanology, USSR Academy of Sciences, Moscow, USSR
(Plate 42; 1 Text figure)
THE CEPHALOPOD FAUNA of the Indian Ocean is by far
not as well investigated as that of the Pacific and Atlantic
Oceans. At present there are nearly half as many species
of oceanic squids (Oegopsida) known in the Indian Ocean
as in the Atlantic. From the general zoogeographic con-
ception of the richness and diversity of the Indo-West-Pa-
cific fauna a contrary relation might be supposed. The
number of species of neritic squids (Myopsida) for instance,
which can be caught easier than Oegopsida and so could »
be better studied, is by a third as large in the Indian Ocean
as in the Atlantic. Now the investigations of the Indian
Ocean are intensified and so our knowledge of its fauna,
and its cephalopods in particular, has increased.
Two specimens of the rare squid Chaunoteuthis mollis
ApPELLOF, 1891, both in excellent condition, have been
found in a small collection of pelagic cephalopods from
stomachs of handsawfishes (Alepisaurus spp.), captured
by Mr. V. G. Osipov of the Pacific Research Institute of
Fisheries and Oceanography, Vladivostok, in the tropical
eastern Indian Ocean aboard the R/V Orlik. So far, this
species has been found only in the Atlantic Ocean and the
Mediterranean Sea. A review of the collection and some
considerations about the feeding of Alepisaurus and the
importance of cephalopods in their food are given in an-
other article (Parin, Nesis & Vinocrapov, 1969). The
present paper will deal only with the description of the
Chaunoteuthis mollis found.
I owe thanks to Dr. N. V. Parin for making the material
available, and to Dr. Johanne Kjennerud of the Zoologi-
cal Museum, Bergen, for the re-investigation of Appellof’s
type specimen in that Museum and for sending me valu-
able information.
TEUTHIDA
Oegopsida
ONYCHOTEUTHDAE
Chaunoteuthis mollis APPELLOF, 1891
From the stomach of an Alepisaurus sp.: 13°00’S, 101°26’
E, November 25, 1963, two specimens, both females. Dor-
sal mantle length (DML) of no, 1: 121 mm, of no. 2:
125 mm; total length 194mm and 204 mm, respectively.
Weight of both specimens together: 40.4 g (Plate 42, Fig-
ure 1).
The body is very soft, flabby, the skin thin, but rather
firm. The mantle muscles are of gelatinous consistency.
The mantle form is conical; it is widest at the anterior
margin: mantle width = 45 to 46% DML. The anterior
margin of the mantle shows a small blunt projection on
the dorsal side and hardly visible lateral projections. The
ventral margin is very slightly emarginated, nearly straight.
The ventral mantle length is 96 to 98% DML. The gladius
is not visible through the mantle, but there is a narrow
dorsal groove over the gladius between the fins.
The fins are transversely-rhombic; their length is 46 to
49% DML, the width is 96% DML.
The head is a little narrower than the mantle, flattened
dorsally; its width is 17 to 19% DML. There are some 8
nuchal folds on either side.
The eyes are big; their length is 114 to 12% DML; the
width is 10 to 104% DML. The eyelid is circular, with a
shallow sinus anteriorly. One luminous organ, long and
band-like, with slightly widened edges and a narrowed
middle part is situated on the ventral periphery of the
THE VELIGER, Vol.
12, No. 3
[Nests] Plate 42
Figure 1 Figure 2
Figure 1: Chaunoteuthis mollis APPELLOF. Specimen no I
Two-thirds natural size.
Figure 2: Chaunoteuthis mollis AppeLOr. Eye. An ocular photo-
phore is seen. Three times natural size.
Vol. 12; No. 3
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Page 291
eye-ball. The organ’s colour is brightly orange. Its dimen-
Sions in) no! — 1/2) <3)mm) in) no! 27—" 14<3)mm
(Plate 42, Figure 2).
The funnel extends almost to the vertical of the anterior
eye margin. The funnel groove is triangular, posteriorly
restricted by a softly arched skin ridge.
The funnel cartilages are simple, lengthened, slightly
broader posteriorly, with a narrow, deep, very slightly
S-shaped groove. The mantle cartilages are simple thin
ridges nearly twice as long as the grooves of the funnel
cartilages.
The arms are nearly equal in length, their order is
2°3°4-1. The length of the dorsal arms is 31 to 36%
DML, of the dorso-lateral it is 403 to 464%; that of the
ventro-lateral arms is 38 to 45% and that of the ventral
arms is 34 to 40% DML. All arms have swimming mem-
branes, which are narrow at the dorsal arms, wider at the
dorso-lateral ones and rather wide at the ventro-lateral
and ventral arms. Protective membranes are nearly in-
conspicuous. The suckers are small, their horny rings
without dentition, and callose distally. The ring callus is
so great that the space of the ring looks excentric or like
the pupil of an octopus or a cuttlefish, and sometimes is
even closed totally. The tentacles are reduced to only
short scraps of stems; their length is 5 to 64% DML.
The gladius is of a deep brown colour, narrow, strongly
convex, especially in its posterior half; the vane is slightly
widened, the lateral edges are not thickened, transparent.
The colour of the animal is a pale brown above and an
even more pale brown below. Large and small brown
chromatophores are dispersed so that a characteristic eye-
spot pattern arises, especially in the posterior part of the
mantle and on the dorsal side of the fins. The ventral side
of the fins (except for the edges) and of the first 3 pairs
of the arms, and the dorsal side of the ventral arms are
not coloured at all. No “very fine scales,” that have been
seen in this species by CLarKE & Maui (1962) were
found in the present specimens.
The stripes of spermatophores — typical for this spe-
cies and almost unique among cephalopods — are found
in both females. They are located at one side of the vent-
ral mantle surface, in the area of the mantle-locking
cartilages. Both females have only one such stripe: no. 1
at the left and no. 2 at the right. The length of the stripes
is 35 mm and 27 mm, respectively; they start 4mm from
the anterior edge of the mantle. The number of spermato-
phores is 26 and 36, respectively; the spermatophores are
attached to the outer surface of the mantle in a narrow
and shallow groove that looks like a closed cicatrice. The
shape of the spermatophores is similar to that of a sipun-
culid worm of the genus Golfingia, the “bodies” of the
spermatophores, that is the sperm reservoirs, are firmly
immersed in the tissues of the mantle, the “heads” and
“necks” protrude outward.
The eggs of both females are not much developed; their
size is 0.35 to 0.40 mm.
DISCUSSION
At present there are 7 specimens of Chaunoteuthis mollis
known: 6 adults (ApPELLOF, 1891; LONNBERG, 1896;
PrerFer, 1912; Jousrn, 1920; Grimpz, 1921, and
Torcuio, 1967) and one young with a mantle length of
1.25 cm (Nasr, 1923). All the adults were found on the
surface of the Mediterranean Sea and in the Eastern
Atlantic Ocean (Text figure 1). The only young specimen
was caught also in the Mediterranean Sea at a depth of
150 m.
The specimens investigated are in accord with the de-
scriptions of preceding authors except for the eye photo-
phores, the absence of which is mentioned even in the
generic diagnosis (PFEFFER, 1912). At my request Dr.
Johanne Kjennerud re-investigated Appelof’s type speci-
men kept in the Bergen Museum and was so kind to
inform me that on the only eye-ball of this specimen there
is a structure which may be a luminous organ. It consists
of a band of very fine dark chromatophores with a layer
behind, which seems to be iridescent.
As for their shape and position the luminous organs of
Chaunoteuthis mollis are very like the organs of Ctenop-
teryx siculus (family CTENOPTERYGIDAE).
All 8 now known adult specimens of Chaunoteuthis mol-
lis are females, and 7 of them have had stripes of sperma-
tophores on the outer surface of the mantle: 4 on both
sides of the median line, 3 on one side only. The number
of spermatophores in one female may reach 100 (Tor-
cHIo, 1967). The stripes of spermatophores start always
a little off the anterior edge of the mantle and extend
nearly parallel to the median axis of the body. But their
length may differ; they may diverge or fork. Probably
they are produced at copulation, once in the life, perhaps
even in the youth of the female. Grimpr (1921) and
Torcuio (1967) speculated that the male cuts the skin of
the female with his beak and places the spermatophores
along the cut. Then the edges of the wound close and the
spermatophores remain firmly fixed in the tissues of the
female. This speculation seems to be correct.
Chaunoteuthis mollis was considered a deep-water
species (GrimpE, 1921). Probably this is not so. The
hooks of the long line, by which the Alepisaurus that had
swallowed the Ch. mollis were caught, were at a depth
Page 292
THE VELIGER
Vol. 12; No. 3
Figure 1
Distribution of Chaunoteuthis mollis
(@ = known records; © = new record)
of 80-90 to 120-150 meters. The main habitat of Alepi-
Saurus is in the lower epipelagic and upper mesopelagic
(Parin, 1968; Parin, Nests « ViNocRADOV, 1969). It is
most probable that Ch. mollis inhabit the lower part of
the surface zone (epipelagic) and the transitional zone
(mesopelagic).
SUMMARY
An onychoteuthid squid, Chaunoteuthis mollis, is recorded
for the first time in the tropical eastern Indian Ocean: 2
females were found in the stomach of an Alepisaurus sp.
An ocular photophore, previously unknown, is described.
It is supposed that Ch. mollis inhabit the lower epipelagic
and mesopelagic zones.
LITERATURE CITED
APPELLOF, A.
1891. Teuthologische Beitrage, II. Chaunoteuthis n. g. Oegop-
sidarum. Bergens Mus. Aarsber. for 1890, no. 1: 1-29
Ciarxe, M. R. « G. E. Maur
1962. _A description of the “scaled” squid, Lepidoteuthis grim-
aldi Jounin, 1895. Proc. Zool. Soc. London 139 (1):
97 - 118
GrimPE, GEorc
1921. Teuthologische Mitteilungen, VI. Das Leipziger Stiick
von Chaunoteuthis mollis ApPpELLOF. Zool. Anz. 52 (12/13):
289 - 296
Jousin, Lours
1920. Céphalopodes provenant des campagnes de la “Prin-
cesse Alice” (1893-1910). (3° série). Rés. Camp. Sci. Monaco
54: 1-113; 14 plts.
LONNBERG, E.
1896. Notes on some rare cephalopods.
skaps-Akad. Férhandl. 8: 603 - 612
Narr, ADOLF
1923. Die Cephalopoden. Systematik. Fauna u. Flora Neapel
Monogr. 35, 1 (1), no. 2: i-xiv+149 - 863; 473 figs.
Parin, N. V.
1968. Ichthyofauna of the oceanic epipelagial.
Moscow: 1 - 186 (in Russian)
Parin, N. V, K. N. Nests « M. E. Vinocrapov
1969. Materials on the feeding of the Alepisaurus in the Indian
Ocean. Voprosy Ikhthiologii 9 (3/561) : 526 - 538 (in Rus-
sian)
PFEFFER, GEorc J.
1912. Die Cephalopoden der Plankton-Expedition. Zugleich
eine monographische Ubersicht der Oegopsiden Cephalopoden.
Ergsb. Plankton-Exped. Humboldt Stift. 2: i-xxi+1-815;
Atlas 48 plts.
Torcuio, M.
1967. Eccezionale reperto di Chaunoteuthis mollis (ApPEL-
LOF) nello Stretto di Messina. Natura (Milano) 58 (3):
193 - 207
Ofv. Kgl. Veten-
“Nauka”,
Vol. 12; No. 3
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Page 293
A Note on the Chromosome Number and Interrelationships
in the Marine Gastropod Genus Thais
of the United States Pacific Coast '
BY
MUZAMMIL AHMED’
AND
ALBERT K. SPARKS
College of Fisheries, University of Washington, Seattle, Washington 98105
(Plate 43)
SEVERAL SPECIES OF MARINE SNAILS belonging to the
family Muricidae (Gastropoda : Streptoneura) inhabit
the Pacific Coast of North America. The genus Thais is
represented by 4 species, namely, T: lamellosa (GMELIN,
1791), T. lima (GmeEttn, 1791), T: emarginata (DESHAYES,
1839), and T: canaliculata (Ductos, 1832). Although these
are customarily recognized as four distinct species, K1n-
carp (1957, 1964) considers the last three as only “morphs”
of a single polytypic species, T: lima. Thais lamellosa and
T. canaliculata are easily distinguished but specimens of
T. lima and T. emarginata show intergradation in shell
morphology and are difficult to tell apart.
The West Coast species and morphs of Thais are ex-
tremely abundant on exposed and sheltered rocky coasts.
They display a wide variety of form, sculpture and color.
A pelagic larval stage is absent and the young animals
emerge from egg capsules which are deposited in large
numbers by the snails almost throughout the year. The
dispersal of these snails would seem to be restricted to the
immediate vicinity where they emerge from egg capsules.
The presence of mud or sand around rocks and increasing
depths constitute barriers for their migration. As a conse-
quence “microgeographical” populations are established
on different islands and even on isolated stretches of
* Contribution No. 313 from the College of Fisheries, University
of Washington, Seattle, Washington 98105
2 Permanent address: Department of Zoology, University of Kar-
achi, Karachi, Pakistan
land on the same beach. The conditions for genetical
differentiation are highly favorable in such populations.
We have undertaken an investigation of the chromo-
somes of the above named species of Thais from the West
Coast of the United States in order to elucidate the cyto-
genetic structure of their microgeographical populations.
A previous study of similar populations of the European
T. lapillus (Linnakzus, 1758) by Statcer (1954, 1957)
indicated the existence of numerical and structural chro-
mosome polymorphism. The extreme forms of this species
occurring on exposed and sheltered beaches possessed
haploid numbers of 13 and 18 respectively and forms
present on intermediate wave exposed beaches had inter-
mediate numbers. Centric fusions due to reciprocal trans-
locations were shown to be responsible for the Robertsonian
nature of the chromosome polymorphism. NisHrKAWA
(1962) examined 3 species of Thais, T: bronni (DUNKER,
1860), T: clavigera (Kiister, 1858), and T: luteostoma
(Hotten, 1802) from Japan and found 30 to be the
common haploid number. He did not find any numerical
chromosome polymorphism.
We have examined chromosomes from oocytes obtained
from egg capsules. The egg capsules of the 4 species were
collected from the coastline of the States of Washington,
Oregon and California. The standard aceto-orcein squash
technique of chromosome preparations was routinely used.
Although Thais lima and T. emarginata capsules contain
nurse eggs, 7: lamellosa does not and the shape and size
of its egg capsules and the number and size of its oocytes
Page 294
also differ from the other two. We found that T: canalicu-
lata egg capsules and oocytes possess a greater resemblance
to those of T: Jamellosa rather than to those of T: lima
and T: emarginata.
A haploid chromosome number of 35 is common to
the 4 species. Three other marine gastropod species,
Ceratostoma (Purpura) foliatum (Gmeuin, 1791), Oci-
nebra japonica (DUNKER, 1860) and Fusitriton oregon-
ensis (REDFIELD, 1848), which were examined by us also
share this number. In prometaphase and metaphase-I
plates 35 bivalents were counted with clarity. In all cases
a definite prometaphase stretching of bivalents was ob-
servable (Plate 43, Figures 1 and 2) as in Thais lapillus
(StaicEr, 1954). In the 4 species of Thais, the prometa-
phase bivalents may stretch as long as 40m or more.
Bivalents were generally rod-like, formed by acrocentric
(subterminal) or telocentric (terminal) chromosomes and
possessed a single chiasma at prometaphase or metaphase-
I. Associations of meiotic bivalents, such as quadrivalents
and, in some cases, linear trivalents, were observed so that
meiotic plates were polymorphic for interchanges or
translocations. In a total of 100 animals or more the
chromosome number was usually N = 35.
In mitotic plates of the 4 species 70 chromosomes were
usually seen. The majority of chromosomes were acrocent-
ric or telocentric but metacentrics were present. The
chromosome number is very large and difficult to work
with. A detailed study of the meiotic and mitotic chromo-
somes of several populations is being made despite this
difficulty.
This study of egg capsules, oocytes, and chromosomes
of several populations of 'Thais forms and species throws
THE VELIGER
Vol. 12; No. 3
some light on the confused status of their taxonomy and
interrelationships. We are inclined to conclude tentatively
that while T: lima and T. emarginata may be the morphs
or forms of the same species, T: lamellosa and T. canalt-
culata are distinct species and perhaps closer to each other
than to T. lima and T. emarginata. Kincam’s suggestion
(1964) that T: lima, T: emarginata and T: canaliculata be
consolidated under one species may be valid only for the
first two of this series. Thais canaliculata shows consider-
able differences in shell morphology, shape of egg capsules
and number and size of oocytes.
LITERATURE CITED
Kincaip, TREVOR
1957. Local races and clines in the marine gastropod Thais
lamellosa (GmEuIN). A population study. Calliostoma Press
1964. Notes on Thais (Nucella) lima (GMELIN), a marine
gastropod inhabiting areas in the North Pacific Ocean.
Calliostoma Company, Seattle.
NISHIKAWA, SHYOHEI
1962. A comparative study of chromosomes in marine gastro-
pods with some remarks on cytotaxonomy and phylogeny.
Journ. Shimonoskei Coll. Fish. 11 (3): 149 - 186
STAIGER, HANSRUDOLF
1954. | Der Chromosomendimorphismus beim Prosobranchier
Purpura lapillus in Beziehung zur Okologie der Art. Chro-
mosoma 6: 419 - 478
1957. | Genetical and morphological variation in P lapillus
with respect to local and regional differentiation of population
groups. Coll. Intern. Biol. Mar. St., Roscoff; L’Année Biol.
3: 33; fasc. 5-6: 252 - 258
THE VELIGER, Vol. 12, No. 3 [AHMED & Sparks] Plate 43
Figure 1
Meiotic bivalents in a prometaphase plate of Thais emarginata.
Approximately X 1000. Phase contrast.
Figure 2
Camera lucida drawing of prometaphase meiotic bivalents
of Thais emarginata.
vi
0h
Vol. 12; No. 3 THE VELIGER
Page 295
Five New Species of Terebra from the Eastern Pacific
BY
TWILA BRATCHER
8121 Mullholland Terrace, Hollywood, California 90046
AND
R. D. BURCH
P.O. Box 133, Downey, California 90241
(Plate 44)
IN EXAMINING the numerous specimens of West American Shy — Laura Shy Collection, Westminster,
Terebridae made available to us during the past two years, California
we have encountered several species which have not been SU — The Conchological Collection of Stanford
previously described. These species have been seen with University
some frequency in several of the larger institutional col- Thomas -—- Lawrence Thomas Collection, Morro Bay,
lections as well as in private collections. California
Terebra species from West America, in several instances, .USNM -— United States National Museum
exhibit variations in shell characteristics which make iden- YPM — Yale University Peabody Museum of
tification very difficult unless a long series of comparison
specimens is available, and good protoconchs are a desir-
able aid in separating species which have somewhat simi-
lar shell characteristics.
Abbreviations have been used for a number of the
institutional collections cited here; these, along with the
location of the private collections cited, are as follows:
AHF — Allan Hancock Foundation (Material on
loan to the Los Angeles County Muse-
um of Natural History)
AMNH~ - American Museum of Natural History
ANSP — Academy of Natural Sciences of
Philadelphia
B&B — Bratcher and Burch Collection, Los Angeles,
California
BM(NH) -— British Museum (Natural History)
CAS — California Academy of Sciences
DMNH_ -— Delaware Museum of Natural History
Frisbey | — Jeanne Frisbey Collection, Port Isabell,
Texas
LACM __— Los Angeles County Museum of Natural
History
SBMNH - Santa Barbara Museum of Natural History
SDMNH - San Diego Museum of Natural History
Natural History
Terebra shyana BRATCHER & BURCH, spec. nov.
(Plate 44, Figures 9, 10)
Description: Size medium, slender; color dark buff with
darker fulvous blotches; whorls slightly convex with noded
subsutural band set off by well marked suture and sub-
sutural groove; nucleus of 3 glassy slender whorls; early
sculpture of numerous slightly curved axial ribs which are
equal to their interspaces and of evenly spaced spiral
cords (4 on 3™ postnuclear whorl) which cross the ribs
faintly; ribs continuous from suture to suture with sub-
sutural groove forming elongate nodes on band; in later
whorls axial ribs become slightly narrower than inter-
spaces, spiral cords crossing ribs form slight nodes giving
a cancellate appearance, and spiral cords are more nu-
merous (6 on penultimate whorl) and less evenly
spaced; body whorl of average length with noded axial
ribs continuing to periphery and fading out posterior to
keel of siphonal fasciole; aperture semi-elongate; outer
lip thin with color blotches showing through; columella
curved with a slight plication, moderately laminated;
siphonal fasciole striate with sharp posterior keel; anterior
Page 296
canal broad and long; length 26.0 mm; diameter 5.6 mm;
14 whorls plus nucleus.
Holotype: LACM, Type Collection no. 1249.
Type locality: All specimens of the type material were
collected at Manzanillo, Colima, Mexico, 19°02’ N Lat.,
104°21’W Long.; 17-40 fathoms; by Laura Shy and
Jeanne Frisbey.
Paratypes: Paratypes are deposited in the following col-
lections: CAS, Department of Geology, Type Collection
no. 13280; SU Conchological Collection no. 9992; USNM
no. 679481; Shy; Frisbey;B & B no. 742.
Other material examined: In addition to two lots of type
material, we have examined 9 lots of this species: CAS no.
27581 from near Santa Isabel Island, Gulf of California;
AHF no. 1031-40 from Santa Maria Bay, outer coast of
Baja California; AHF no. 765-38 from Chacaua Bay,
Mexico, at 5 - 10 fathoms; AHF no. 1753-49 from north
of Coyote Point, Gulf of California; AHF no. 273-34
from Tenacatita Bay, Mexico, at 45 fathoms; AMNH
no. 88725 from the Gorda Banks, Gulf of California;
AMNH nos. 77796 and 77826 from Coronados Islands,
Gulf of California, Mexico.
Largest specimen examined: length 36.1 mm; diameter
7.5 mm.
Discussion: Specimens vary in the number of axial ribs
and spiral cords. One of the largest specimens examined
has ribs much narrower than the interspaces and only 4
rows of unevenly spaced spiral cords which cross the ribs
without producing a cancellate appearance. The largest
specimen appears cancellate from the first postnuclear
whorl through the body whorl.
Terebra shyana resembles several other Terebra species
which occur in the Indo-Pacific and Panamic regions. It
can be separated from T: panamensis Datu, 1908, by the
latter’s heavier, more cancellate sculpture and less conspic-
uous subsutural band, while T: turrita E. A. Smiru, 1873,
of Australia, T. serotina Apams & ReeEve, from Ja-
pan and the Philippine Islands, and T. textilis Hinps,
1844, of the Indo-Pacific, may be distinguished from T.
shyana by their more cancellate sculpture, flatter whorls,
protoconchs and style of columella. Another Panamic
Terebra, the description of which is presently in press, has
a rather close superficial resemblance to T: shyana, there-
fore a close examination of this species is desirable, espe-
cially of the protoconch and of the spiral sculpture
anterior to the periphery of the body whorl.
This species is named in honor of Mr. and Mrs. Carl
Shy of Westminster, California, for their having collected
the first specimens of this species which came to our atten-
THE VELIGER
Vol. 12; No. 3
tion and for their generosity in having made their large
collection of specimens available for study.
Terebra brandi BRATCHER & BURCH, spec. nov.
(Plate 44, Figures 5, 6)
Description: Size medium small, slender; color bluish
gray; whorls slightly convex with moderately convex sub-
sutural band marked by impressed suture and subsutural
groove which crosses axial ribs after the 4° postnuclear
whorl; nucleus of 4 dark brown, shiny, smooth whorls;
sculpture consistent throughout except that subsutural
band becomes discernible after 4 postnuclear whorl;
axial sculpture of slightly curved ribs narrower than
interspaces on both whorl and band; spiral sculpture of
rows of evenly spaced spiral grooves which do not cross
axial ribs, 6 on penultimate whorl, 3 on subsutural band;
body whorl of medium length with axial ribs ending at
periphery and interspaces divided by 7 rows of spiral
grooves; anterior to periphery 7 unevenly spaced spiral
grooves extend to siphonal fasciole; aperture moderately
elongate; outer lip thin; columella brown with slight
curve, very faint plication, light lamination; siphonal fas-
ciole striated with posterior keel; anterior canal moder-
ately long, curved, medium width; length 16.9 mm, di-
ameter 3.9 mm; 10 whorls plus nucleus.
Holotype: LACM —- AHF, Type Collection no. 1252.
Type locality: Petatlan Bay, Mexico, 17°31’N Lat.,
101°27’ W Long.; Allan Hancock Pacific Expedition col-
lecting station 265-34; 5-10 fathoms on hard sand and
shell bottom.
Paratypes: One paratype; LACM - AHF, Type Collec-
tion no. 1253; very poor conditicn.
Other material examined: In addition to the type lot,
AHF 702-37 from Angeles Bay, Baja California, Mexico,
at 18 fathoms; Shy Collection from Manzanillo, Mexico,
7-8 fathoms; AHF 584-26 from Conception Bay, Baja
California, Mexico, intertidal; Frisbey Collection from
Manzanillo, Mexico, 15-40 fathoms; LACM Hill Collec-
tion from Bay of Dulce, Mexico; AHF 763-38 from Cape
Corrientos, Mexico, 5 - 10 fathoms; LACM no. B-16 from
Taboga Island, Panama, 5 fathoms; USNM 566896 from
Mazatlan, Mexico; USNM 426749 from Peru; B « B no.
273 from Panama Bay, Panama, intertidal.
Except for the type lot and one other, each of which
contains 2 specimens, all lots examined were of one speci-
men each.
Largest specimen examined: USNM 566896; length
24.1 mm; diameter 4.6 mm.
Vol. 12; No. 3
Discussion: The variation exhibited among the specimens
examined is confined to the color, which may be lavender
gray, bluish gray or beige gray.
Several species of Panamic Terebra somewhat resemble
T. brandi in sculpture. Terebra berryi CAMPBELL, 1961,
has a larger apical angle, more convex whorls with a
depressed subsutural band, a more pronounced and num-
erous spiral sculpture and different style of color and
color pattern; while 7: churea CAMPBELL, 1964, has a
shorter protoconch, more pronounced subsutural band and
more numerous spiral sculpture than T: brandi. The
western Atlantic Ocean and Gulf of Mexico species, T.
protexta (Conrap, 1848), and T: glossema SCHWENGEL,
1940 and 1942, have some sculptural resemblance but may
be separated by their axial ribs which continue anterior
to the periphery of the body whorl and by their columellas
which are more curved or twisted than the slightly curved
columella of T: brandi.
This species is named for Dr. Louis M. Brand of
Houston, Texas, in recognition of his generous assistance
and early encouragement of our study of the Terebridae.
Terebra dorothyae BRATCHER & BURCH, spec. nov.
(Plate 44, Figures 7, 8)
Description: Size medium; color brown, with portions
of subsutural band and nodes slightly lighter; early whorls
somewhat convex and later whorls almost flat, with mod-
erately convex subsutural band marked by impressed
suture and broad shallow subsutural groove; nucleus
partially missing; first postnuclear whorl glassy with
slightly curved axial ribs narrower than interspaces; next
4 postnuclear whorls with ribs extending from node on
subsutural band to following suture; in later whorls, nodes
on band become elongate with ribs on remainder of
whorl broken into nodes by spiral grooves, 5 on penulti-
mate whorl; body whorl long, with 2 rows of spiral nodes
wide-set and 3 rows, smaller and closer together, posterior
to periphery; anterior to lighter band of color at peri-
phery, 3 rows of cords form small nodes where they cross
axial ribs, followed by 3 rows of cords which do not form
nodes; aperture elongate; outer lip thin with light peri-
pheral band and pattern of nodes showing through; co-
lumella slightly curved with no plication; siphonal fasciole
heavy, striated, with exceptionally large posterior keel;
anterior canal broad; length 29.9 mm; diameter 6.5 mm;
11 whorls plus portion of nucleus.
Holotype: LACM - AHF, Type Collection no. 1250.
Type locality: AHF Pacific Expedition collecting stations
THE VELIGER
Page 297
770-38 and 929-39; near San Jose, Guatemala; 13°53’ N
Lat., 91°09’ W Long.
Paratypes: LACM - AHF, Type Collection no. 1251;
CAS, Department of Geology, Type Collection no. 13281;
SU Conchological Collection no. 9993; SBMNH, Type
Collection no. 27141; SDMNH, Type Collection no. 51249;
AMNH no. 154674; USNM no. 679482; YPM no. 12-
15635; BM(NH) ; DMNH no. 22421; ANSP no. 316223.
Other material examined: AHF 762-28 from inner Gorda
Bank, Gulf of California, Mexico, 60 fathoms; CAS 35006
from coast of Michoacan, Mexico; Shy Collection from
Santiago Bay, Manzanillo, Mexico, 7 - 8 fathoms; Frisbey
Collection from Manzanillo, Mexico, 15-40 fathoms; LA
CM Hill Collection from Panama Bay, Panama; LACM
no. A5498 from the Galapagos Islands.
Largest specimen examined: LACM no. A5498; length
36.0 mm, diameter 7.7 mm; nucleus missing.
Discussion: This species exhibits little variation among
the specimens examined. In some specimens the light color
band at the periphery of the body whorl and the lighter
subsutural band are very distinct while in other specimens
these features are almost non-existent. In occasional spe-
cimens the axial ribs and spiral cords crossing them are
conspicuous, while in others only the nodes are evident.
Terebra dorothyae has little resemblance to other spe-
cies of Terebra; however, some variations of other species
should be considered in making identifications. The Japan-
ese species, T: pustulosa E. A. Smitu, 1879 (= T. granu-
losa E. A. Smiru, 1873, not Lamarck, 1822) has a weak-
ly but broadly impressed subsutural groove and fewer
rows of nodes than are found in T: dorothyae. Of the Pan-
amic species which exhibit rows of spiral nodes, T: tuber-
culosa Hinps, 1844, is a broader and heavier shell with
more conspicuous subsutural band and fewer rows of
spiral nodes, while T: cracilenta Li, 1930, also has a
broader apical angle and heavier shell with more pro-
nounced subsutural band and nodulous sculpture than
T. dorothyae. Both T: ropert Prtspry & Lowe, 1932, and
T. adairensis CAMPBELL, 1964, are slender species, as is
T. dorothyae, but each of these has only 2 rows of nodes
on each whorl, one posterior and one anterior to the
suture. Terebra glauca Hinps, 1844, might also be com-
pared, although this species is consistently broader, less
uniformly noded or colored, and the columella more slan-
ted than is found in specimens of T: dorothyae.
This species is named in honor of Mrs. Dorothy Brown
of San Diego, California, in recognition of her generosity
in making specimens available for study and of her inter-
est in conchology.
Page 298
Terebra allyni BraTCHER & BURCH, spec. nov.
(Plate 44, Figures 1, 2, 3, 4)
Description: Size medium; color beige mottled with rust
brown; whorls flat with slightly convex subsutural band
set off by suture and shallow subsutural groove; nucleus
of 14 dome shaped glassy whorls; first 2 postnuclear
whorls translucent with almost straight ribs about equal
to interspaces and no noticeable subsutural band or spiral
sculpture; in next 6 whorls ribs become more widely
spaced, starting as small elongate nodes on subsutural
band, and wide but feeble spiral cords develop which do
not cross ribs; after 6 postnuclear whorl wide spiral
cords, though inconspicuous, cross ribs forming rounded
nodes, 4 rows on penultimate whorl; fine axial striae
form between rows of nodes and cross subsutural band;
body whorl of average length with 4 rows of nodes, the
anterior being at the periphery, followed anteriorly by
a broad shallow groove and 4 rows of smaller nodes;
anterior to periphery spiral sculpture becomes finer, more
numerous, and continues to keel of siphonal fasciole as
do axial striae; aperture elongate; outer lip sturdy; colu-
mella straight with no plication, thinly laminated; siphon-
al fasciole striated with posterior keel lacking in strength; —
outer canal straight, broad; length 25.8mm; diameter
6.3 mm; 12 whorls plus nucleus.
Holotype: CAS, Department of Geology, Type Collection
no. 13278.
Type locality: CAS station 23779, east shore of Maria
Madre Island, Tres Marias Group, 21°35’ N Lat., 106°26’
W Long.; 5 to 10 fathoms.
Paratypes: CAS Type Collection no. 13279; LACM Type
Collection no. 1254; SBMNH Type Collection no. 27142;
SU Conchological Collection no. 9995; SDMNH Type
Collection no. 51248; USNM no. 679534; AMNH no.
154675; ANSP no. 316224; DMNH Type Collection no.
22421; BM(NH) ; B&B Collection no. 743.
THE VELIGER
Vol. 12; No. 3
Other material examined: In addition to the type lot of
61 specimens, many of which are immature, we have
examined 2 specimens, CAS 29894, Margarita Island,
Lower California, Mexico; 3 specimens, CAS 23810, Es-
piritu Santo Island, Gulf of California; 1 specimen AM
NH 74171, Maria Madre Island, Tres Marias Group, Pu-
ritan Expedition; 1 specimen, AMNH, San Juanito Is-
land, Tres Marias Group, Puritan Expedition; 1 specimen,
LACM 66-8, Santa Margarita Island, intertidal to 6
feet; 15 specimens, LACM 65-16, Banderas Bay, Jalisco,
Mexico, 10 - 15 fathoms; USNM 564817, Santa Inez Bay,
Gulf of California.
Largest specimen examined: LACM 65-16; length 39.1
mm; diameter 8.8 mm.
Discussion: The smaller lots of specimens examined,
which contain 1 or 2 specimens each, have only 1 row
of spiral nodes at the periphery of the body whorl (Plate
44, Figures 3 and 4). These sharply rounded nodes are
formed on axial ribs which fade completely at the poster-
ior as well as the anterior portion of the whorl. This
variation in sculpture is the only form we have seen from
several of the collecting stations, although occasionally it
is found among specimens which contain more sculpture.
The rows of spiral nodes vary from 0 in some specimens to
4 in others examined. Occasional individual specimens are
more slender than others, and some have a more convex
subsutural band with larger nodes. The color varies little
among specimens examined, with the exception of lot LA
CM 65-16 which is light grayish brown with rust colored
blotches between the nodes of the subsutural band.
Several species of Indo-Pacific and Panamic Terebra
have a superficial resemblance to T: allyni with T: specil-
lata Hinps, 1844, resembling the more sculptured form.
Terebra specillata has a more cancellate sculpture with
more prominent subsutural band and consistent heavy
sculpture through all whorls than T: allyni. The less sculp-
tured forms of T. allyni resemble T: conspersa Hinps,
1844, but the latter has numerous incised interstitial spiral
Explanation of Plate 44
Figure 1: Terebra allyni BRATCHER & BurRcH, spec. nov. Holotype
CAS no. 13278 X 34
Figure 2: Terebra allyni. Hypotype, CAS no. 13279; nucleus
Figure 3: Terebra allyni. Hypotype, CAS no. 13279; variation of
sculpture xX 3
Figure 4: Terebra allyni, dorsal view of same shell as in Figure 3
Figure 5: Terebra brandi BratcHER & Burcu, spec. nov. Holotype
LACM - AHF no. 1252 : X 34
Figure 6: Yerebra brandi, nucleus of same shell as in Figure 5
Figure 7: Terebra dorothyae BRATCHER & Burcu, spec. noy. Holo-
type LACM - AHF no. 1250 X 3
Figure 8: Terebra dorothyae. Paratype LACM - AHF no. 1251
Nucleus
Figure 9: Terebra shyana BraTCcHER & BurcH, spec. nov. Holotype
LACM - AHF no. 1249 XK 3
Figure 10: Terebra shyana, nucleus of same shell as in Figure 9
Figure 11: Terebra hancocki BRATCHER & BuRcH, spec. nov. Holo-
type LACM - AHF no. 1255 X14
Figure 12: Terebra hancocki. Hypotype LACM no. 65-23; nucleus
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Vol. 12; No. 3
striations instead of a spiral sculpture of raised cords.
Terebra interstincta Hinps, 1844, has a more slanted
columella, quadrate aperture, enlarged nodes on subsu-
tural band and small nodes posterior to the suture. Tereb-
ra dorothyae, new species, is a more slender shell with
smaller and sharper nodes and a more twisted columella
than T: allyni.
This species is named for Mr. Allyn G. Smith of San
Francisco, California, in recognition of his work in Mala-
cology and for his encouragement of and assistance to
workers in this field.
Terebra hancocki BRATCHER & BURCH, spec. nov.
(Plate 44, Figures 11, 12)
Description: Size large; color shiny pale beige back-
ground with irregular blotches of reddish brown; whorls
moderately convex with convex subsutural band marked
by impressed suture and broad, deep subsutural groove;
nucleus missing; early sculpture of almost straight axial
ribs narrower than interspaces and of 3 rows of spiral
grooves which faintly cross ribs; in later sculpture spiral
grooves become unevenly spaced and broad, with strap-
like raised cords betwen, crossing axial ribs to produce
small nodes and give sculpture a file-like appearance;
subsutural band has rounded nodes on early whorls, elon-
gate nodes on later whorls; body whorl of medium length
with lighter stripe and groove at periphery; anterior to
periphery spiral grooves continue to cross now obsolete
axial ribs with somewhat nodose effect; aperture semi-
quadrate; outer lip sturdy, white within; columella white,
twisted, broad, with 2 heavy plications; siphonal fasciole
heavy, striate, with a sharp posterior keel which continues
to posterior plication of columella; anterior canal broad,
very twisted; length 75.3mm; diameter 15.3mm; 15
whorls, apex missing.
Holotype: LACM -—— AHF Type Collection no. 1255.
Type locality: All type material was collected off La
Libertad, Ecuador 2°08’ S Lat., 81°00’ W Long.
Paratypes: LACM — AHF, Type Collection no. 1256;
CAS, Department of Geology, Type Collection no. 13282;
SU Conchological Collection no. 9995; SBMNH, Type
Collection no. 27143; AMNH no. 155886; USNM no.
678614; ANSP no. 316418; DMNH no. 51367; BM(NH).
Other material examined: AMNH 74:75642; AMNH
51:74585 and LACM A875 from the Tres Marias Islands,
Mexico; USNM 590706, Shy Collection and Thomas
Collection from Manzanillo, Colima, Mexico; AHF 265-
THE VELIGER
Page 299
35 from Petatlan Bay, Mexico; USNM A5498 from Gulf
of Tehuantepec, Mexico; LACM A2777 from Corinto
Bay, Nicaragua; AHF 257-34 and 256-34 from Port Cu-
lebra, Costa Rica; AHF 940-39 and 941-39 from the
Gulf of Dulce, Costa Rica; LACM B-12 from Taboga
area, Panama; AHF 239-23 and 413-25 from Port Utria,
Colombia; AHF 399-35 from Salango Island, Ecuador.
Largest specimen examined: The holotype.
Discussion: The colored areas on the pale beige back-
ground may be purplish-gray, gold, reddish-brown or a
combination of all 3 colors. Though we have examined
many specimens of this species, only 1 has been seen with
nucleus intact. The nucleus is of 34 slender, slightly con-
vex whorls. The first 14 whorls of the nucleus are gray in
color and the remaining 2 whorls are light brown.
The 2 plications on the columella of Terebra hancocki
are the most prominent of any west American species we
have examined.
Several Panamic species resemble Terebra hancocki in
general appearance. Terebra variegata Gray, 1834, is a
broader species which does not become as noded or file-
like in appearance and has a protoconch of 24 whorls.
The more cancellate or noded form of T: glauca Hinps,
1844, has a superficial resemblance, but the latter is a
smaller and more slender species with less heavy plications
on the columella and a protoconch of 24 whorls. Some
of the sculptural variations of T. albocincta CARPENTER,
1857, very closely approach T. hancocki in appearance,
but the former is a smaller and more slender species with
a different color pattern, less prominent columellar plica-
tions and different sculpture in the early whorls. Terebra
dislocata (Say, 1822), which occurs on both coasts of the
Americas, is also a smaller and more slender species with
less heavy columellar plications.
This species is named in honor of Captain G. Allan
Hancock for his contributions to Malacology during the
Hancock Pacific Expeditions.
ACKNOWLEDGMENT
We wish to acknowledge with gratitude the assistance
generously given us by a number of individuals and insti-
tutions by having made comparison specimens available,
providing locality information and other data and for
locating or making available needed reference materials.
In addition to Mr. and Mrs. Carl Shy, Mrs. Dorothy
Brown, Mrs. Jeanne Frisbey and Mr. Lawrence Thomas,
whose collections have been cited here, we wish to express
our appreciation for the loan of pertinent comparison
Page 300
specimens to Mr. and Mrs. Ben Purdy of San Diego,
California, and Mr. and Mrs. John Q. Burch of Seal
Beach, California.
A number of institutional collections were also made
available, providing large numbers of comparison speci-
mens with accurate collection and locality information.
We wish to express our appreciation to Dr. John S.
Garth of the Allan Hancock Foundation, Dr. William K.
Emerson and Mr. William Old, Jr., of the American Mu-
seum of Natural History, Dr. Leo G. Hertlein of the
California Academy of Sciences, Dr. James H. McLean
and Mr. Gale Sphon of the Los Angeles County Museum
of Natural History, Dr. Copeland McClintock and Mrs.
Diane Ryerson of the Peabody Museum of Yale Univer-
sity, and Drs. Harald Rehder and Joseph Rosewater of
the United States National Museum for the loan of speci-
mens from their respective institutions.
We are especially appreciative of the patient assistance
given by Mrs. Dorothy Halmos and Miss Mary Ellen
Pippin of the Hancock Library of the University of South-
ern California and of Mrs. Dorothy Martin, Miss Betty
Begun and Mrs. Azile Kokos of the Los Angeles County
Museum Library for locating much of the reference mate-
rial needed for this study. Other reference material has .
generously been made available by Dr. S. Stillman Berry
and Mr. and Mrs. John Q. Burch from their personal
libraries.
LITERATURE CITED
ApaMs, ARTHUR & Lovett Aucustus REEVE
1848-1850. The zoology of the voyage of H. M.S. Samarang,
under the command of Captain Sir Edward Belcher, during the
years 1843-1846. Mollusca, prt. 2: 25 - 44; plts. 10 - 17. London
CampBELL, G. Bruce
1951. Four new Panamic gastropods.
25 - 28; plt. 5
The Veliger 4 (1):
(1 July 1961)
THE VELIGER
Vol. 12; No. 3
1964. New terebrid species from the eastern Pacific (Mollusca:
Gastropoda). The Veliger 6 (3): 132 - 138; plt. 17
(1 January 1964)
CarPENTER, Puitip PEARSALL
1857. Catalogue of the collection of Mazatlan shells in the
British Museum collected by Frederick Reigen. London: Brit-
ish Mus. i - vit+ix - vxi+552 pp. (with preface by J. E. Gray)
(post June 1857). [Warrington ed. with author's preface, pp.
V - viii, publ. simultaneously for distribution with duplicate col-
lections]
Conrab, TrmotHy AsBBotTr
1848. Tertiary fossil shells. Proc. Acad. Nat. Sci. Philadel-
phia 3 (for 1846 and 1847): 19-27; plts. 1, 2
Gray, Joun Epwarp
1834. [Untitled. Terebra] Proc. Zool. Soc. London, prt. 2:
50 - 63 [pp. 50 - 56 — 26 Sept.; 57 - 63 — Nov. 1834]
Hinps, RicHarp BrinsLey
1844. Descriptions of new shells collected during the voyage
of the Sulphur, and in Mr. Cuming’s late visit to the Philip-
pines. Proc. Zool. Soc. London for 1843, pt. 11: 149 - 168
(June 1844)
Li, Coty CHANG
1930. The Miocene and Recent Mollusca of Panama Bay.
Bull. Geol. Soc. China 9 (3): 249-296; plts. 1-8; 1 map
(October 1930)
Pitspry, Henry Aucustus « Hersert N. Lowe
1932. | West Mexican and Central American mollusks collec-
ted by H. N. Lowe 1929-31. Proc. Acad. Nat. Sci. Philadel-
phia 84: 33 - 144; 6 figs.; plts. 1-17; 2 photographs
(21 May 1932)
Say, THoMAS :
1822. An account of some marine shells of the United States.
Journ. Acad. Nat. Sci. Phila. 2 (2): 221-248 (June 1822)
ScHWENGEL, JEAN SANDERSON
1940. [Untitled]. The Nautilus 53 (3): plt. 12 (Jan. ’40)
1942. | New Florida marine mollusks. The Nautilus 56 (2):
62 - 66; pit. 6 (October 1942)
Smrrn, Epcar ALBERT
1873. | Remarks on a few species belonging to the family Tere-
bridae, and descriptions of several new forms in the collection
of the British Museum. Ann. Mag. Nat. Hist., (ser. 4),
1879. Mollusca from Japan. Proc. Zool. Soc. London
11: 262 - 271
(Terebra: pp. 183 - 186; plt. 19) (August 1879)
Vol. 12; No. 3
THE VELIGER
Page 301
Observations on the Anatomy and Biology
of Two California Vermetid Gastropods
BY
MICHAEL G. HADFIELD
Pacific Biomedical Research Center, University of Hawaii, Honolulu, Hawaii 96822
(Plate 45; 4 Text figures)
INTRODUCTION
THE MARINE GASTROPODS included in the family Vermeti-
dae form a morphologically distinct group characterized
by shells which are uncoiled and cemented to a substra-
tum. This feature is shared with the prosobranchs of the
family Siliquariidae to which the Vermetidae are closely
related. The siliquariids are distinct from the vermetids
in having a long slit through the mantle wall and shell on
the right side beneath the rectum, and in retaining a
spiral corkscrew shape in their shell (Morton, 1951,
1955). Both families are placed in the mesogastropod
super-family Cerithiacea, which contains, in addition,
such well known temperate forms as Turritella, Cerithium,
and Bittium (THIELE, 1931; Morton, 1958).
There have been very few comprehensive treatments of
the vermetids. Between 1900 and 1940 only 3 papers
appeared which were devoted to aspects of the biology of
vermetids (though taxonomic citations occur in other
works) ; these were BortTcerR (1930), Yonce (1932),
and Yonce & ItEs (1939). BoETTGER was concerned with
nutritional physiology, while YoncE & ILES compared cer-
tain points of the anatomy of Serpulorbis gigas (Brvona-
Bernarpi, 1832) of the Mediterranean with that of
Dendropoma maximum (SoweErsy, 1835) from Australia.
Since 1940 the vermetids have received considerable
attention at the hands of Professor J. E. Morton of Auck-
land (1950; 1951, a, b, c; 1955; 1965). It was Morton
who separated the vermetids and siliquariids into two
taxa (1951) and attempted to establish sound anatomical
criteria for the generic groupings within the Vermetidae
(1965). Keen (1961) had already performed the ex-
ceedingly difficult task of reducing the more than 45
nominal genera to a realistic 5.
Morton has been particularly interested in mechanisms
of feeding in the vermetids and in the evolution of the
group. While he did have access to preserved material
of Petaloconchus montereyensis (Dati, 1919) and Ser-
pulorbis squamigerus (CARPENTER, 1857), the species
studied here, Morton’s discussion of function in these
species is based upon inferences from anatomy. Morton’s
descriptions of the digestive system are complete, but he
has only briefly described the reproductive system and has
omitted reference to the nervous system altogether.
Compared with that for most other large prosobranch
families, our knowledge of the biology of living ver-
metids is slight, and concerns only a small number of spe-
cies. Of some 250 nominal species of Vermetidae, only
8 or 9 have been examined alive. Morton (1965) dis-
cussed most of the literature on this topic and the only
subsequent work is a study by Urret Sarriet (1966)
on the “vermetid formations” of the Israeli shore of the
Mediterranean Sea in which he presents an interesting
description of the geological effects of vermetid intertidal
zonation.
The following discussion of anatomy and biology of the
two California vermetid species, Petaloconchus monterey-
ensis and Serpulorbis squamigerus, provides a background
for detailed information on reproduction to be discussed
in subsequent papers, and presents certain new information
on systems in these species for which published accounts
are lacking.
MATERIALS ann METHODS
Serpulorbis squamigerus (CARPENTER, 1857)
While a few specimens of this species were collected on
the Monterey Peninsula, most of the animals used were
taken at Malibu and Newport Beach, California. For the
sake of comparison, animals were also collected at San
Diego, La Jolla, Newport Bay, and Santa Barbara, Cali-
Page 302
THE VELIGER
Vol. 12; No. 3
fornia. The animals collected at these localities, though
differing in their habitats, were all clearly of one species.
Petaloconchus montereyensis (DAL, 1919)
Most of the animals utilized in this study were collected
from two restricted populations on Mussel Point adjacent
to the Hopkins Marine Station, Pacific Grove, Califor-
nia. Other specimens, taken for comparative purposes,
were collected at Point Pinos and Pescadero Point, on the
Monterey Peninsula.
Both species were kept for periods of up to more than
one year at 12° to 16° C in aquaria at Hopkins Marine
Station. The aquaria were provided with a continuous
flow of fresh seawater pumped from Monterey Bay.
While the animals survived well in the laboratory, they
usually ceased reproductivity within one to two weeks
after their confinement in aquaria. Except in feeding
experiments, no attempt was made to provide food for
the animals other than the plankton which passes through
the filters at the seawater intake.
The living animals were examined both grossly and
microscopically in small dishes of seawater under a dis-
secting microscope. It was necessary to anesthetize ani-
mals before removing them from their shells for obser-
vation and dissection. Anesthetization was brought about
by immersing the animals in an aqueous solution of mag-
nesium chloride isotonic to sea water (75 g per 1 1 of tap
water). The animals were usually completely immobilized
by this treatment in 3 to 4 hours.
ANATOMY
Serpulorbis squamigerus and Petaloconchus monterey-
ensis, like all the members of the family Vermetidae, con-
struct calcareous shells which are securely cemented to
a substratum. While the configuration of these shells (see
KEEN, 1961) is unlike that of most gastropods, the animals
themselves possess fairly typical gastropodan bodies which
may be divided into head-foot, pallial, and abdominal
regions (Text figure 1; Plate 45, Figures 3, 4).
The foot is cylindrical, naked in Serpulorbis but covered
by an operculum in Petaloconchus (Text figure 4A; Plate
45, Figure 2). The muscular mass of the foot is directly
connected with the well-developed columellar muscle
which runs up the ventral side of the pallial region. Since
the major portion of the adult foot bears the operculum
in most vermetids, this part of the foot is considered to
correspond to the metapodium of the foot of the vermetid
juvenile and other free-living snails (see Morton, 1955).
The mesopodium is represented in the adult vermetid by
a small shield-shaped pad of tissue below the mouth,
Os
Ct
Figure 1
Female Serpulorbis squamigerus with the mantle wall opened along
the dorsal mid-line and deflected to show pallial structures.
Ct - ctenidium; K - kidney; G - ovary; Os - osphradium ;
POd - pallial oviduct R - rectum V - ventricle
while the orifices of the large pedal mucus glands and the
pedal tentacles together represent the propodium. The
two pedal tentacles, which originate in depressions ventro-
lateral to the mouth, are very extensible, and each has a
groove running up its mesial margin.
Vol. 12; No. 3 THE VELIGER Page 303
The snout, with the horizontal slit-shaped mouth at its
terminus, is directly above the propodium. Slightly poste-
rior to the snout, on each side of the dorsal surface of the
head, a cephalic tentacle arises; each bears an eye on its
posterior lateral margin. The head narrows behind the
eyes to form a “neck” region at the entrance to the mantle
cavity.
In life, the head and foot of both species are mostly
black with small amounts of orange pigmentation around
the foot and the lateral ridges of the head. However, the
color pattern of Petaloconchus montereyensis is quite vari-
able, and in some populations the predominant ground
color of the head is brownish-orange. The thickened edge
of the mantle which encircles the head and foot is brown-
ish-orange with flecks of dense white.
The mantle cavity is relatively deep, extending for 50%
of the total length of the animal in Serpulorbis and 40%
in Petaloconchus. The roof of the cavity displays the only
strongly sexually dimorphic character in Serpulorbis.
Here, in reproductively active females, a long slit in the
median dorsal mantle wall provides contact between the
mantle cavity and the shell over the pallial region (Plate
45, Figure 4). The egg capsules are attached to the ex-
posed shell in this region. No such slit occurs in the females
of Petaloconchus.
A ridge of muscular tissue arises on the posterior dor-
sal surface of the head and runs posteriorly, dividing the
ventral portion of the mantle cavity into right and left
halves (Text figure 2A). The ridge flattens out only in the
most posterior portion of the pallial cavity. The organs
lying within the mantle cavity are those of a typical
prosobranch mesogastropod (Text figure 1). They in-
clude an elongate ctenidium on the left wall composed of
triangular lamellae, a long osphradium lying beneath the
gill, a hypobranchial gland on the mantle cavity roof, the
rectum on the right dorsal portion of the roof, pallial re-
productive structures in the posterior right ventral region,
Figure 2
(adjacent column —>)
Diagrammatic cross-sections through Serpulorbis squamigerus
A: through the anterior portion of the mantle cavity of a female;
B: through the posterior portion of the mantle cavity;
C: through the abdominal region of a sexually mature male.
DD - digestive gland duct; COL - columellar muscle;
Cp-capsule gland; Ct-ctenidium; Dg - digestive gland;
E -esophagus; G - gonad; Hy - hypobranchial gland;
K - glandular portion of kidney; MC - mantle cavity;
MG - pedal mucous gland; Pc - pericardium R - rectum
N- right pleuro-visceral nervous connective; VD - vas deferens
Page 304
and the kidney orifice in the posterior wall (Text figures
1, 2A, 2B). The kidney consists of a large sac placed
directly behind the mantle cavity on the right half of the
body, and a large amount of excretory tissue which extends
out into the posterior roof of the mantle cavity (Text
figure 2B). On the left side the pericardium extends dor-
sally and anteriorly over the more posterior part of the
pallial cavity (Text figure 2B). It contains the large ven-
tricle and the smaller auricle of the heart.
The mouth opens into a buccal cavity surrounded by a
massive buccal muscular apparatus which manipulates the
jaws and radula. The buccal cavity opens dorsally into the
esophagus which is provided with a pair of salivary glands
borne on the posterior dorsal surface of the buccal mass.
The-esophagus is very long and runs from the buccal mass
directly posteriorly beneath the mantle cavity, the heart,
and the kidney, to the stomach. The large pedal mucous
glands lie parallel to the esophagus along half its length
(Text figure 2A).
Behind the heart and kidney lies the abdominal region
of the body. The anterior portion of this region contains
several coils of the intestine, the stomach, and the style
sac. Behind the stomach is a tail-like region of variable
length which contains the digestive gland on the left and |
the gonad on the right (Text figure 2C.). The major chan-
nel of the digestive gland is ventral in position, as are the
major blood lacunae, the abdominal nervous extensions,
and the gonoduct.
The elongate columellar muscle originates in the foot,
forms the ventral portion of the body mass in the pallial
region, emerges from the body at the level of the posterior
end of the pallial cavity, and continues backward as a
long free strap to its insertion far back inside the shell.
The nervous system of vermetid gastropods has been
examined only in Vermetus triqueter (BIvoNA-BERNARDI,
1832) by Lacaze-Dutuirrs, 1860. The ganglia and ma-
jor nerves of Serpulorbis squamigerus are easily examined
after anesthetization and formalin fixation of the animal.
The details of the nervous system of this species are pre-
sented in Text figure 3. All major ganglia are paired, a
condition generally accepted as primitive in gastropods
(FRETTER & GRAHAM, 1962), and all except the parietal
(or supra- and sub-esophageal) ganglia lie in their ex-
pected complementary bilateral positions. The sub-eso-
phageal ganglion is located next to the posterior face of
the left pleural ganglion. The supra-esophageal ganglion
is well removed from the circumesophageal nervous com-
plex; it lies more posteriorly and to the left, near the
osphradium.
The cerebral ganglia lie on the esophagus posterior to
the buccal mass. They are provided with thick connectives
to the pleural and pedal ganglia and are linked across the
THE VELIGER
Vol. 12; No. 3
esophagus by a thick cerebral commissure. Three large
nerve trunks leave each cerebral ganglion anteriorly. The
most lateral of these bifurcates, one nerve going to the
region of the mouth and the other recurving and pene-
trating the buccal mass to connect with the buccal gang-
lion on that side. The buccal ganglia lie on the posterior
dorsal face of the buccal mass, and innervate the complex
musculature of the buccal apparatus. The middle nerve
arising from each cerebral ganglion runs to the lips and
the innermost nerve innervates the eye and cephalic
tentacle on its side.
The pleural ganglia lie ventrally to the cerebral ganglia
with which they are connected by thick commissures. A
commissure also links each pleural with the pedal ganglion
of the same side. The left pleural ganglion is closely
associated with the sub-esophageal ganglion. A nerve a-
rises from the dorsal face of each pleural ganglion to
innervate the neck musculature on each side. From the
postero-lateral face of each pleural ganglion a large nerve
arises which receives a zygoneuric connection from the
pleuro-visceral loop. Proximal to the point of zygoneury,
the nerve arising from the left pleural ganglion sends one
branch to innervate the mantle edge on the same side;
distal to the zygoneuric connection from the supra-eso-
phageal ganglion, the nerve runs on to innervate the os-
phradium and ctenidium. The corresponding nerve orig-
inating from the right pleural ganglion receives the zygo-
neuric connection from the sub-esophageal ganglion and
extends toward the right side; it soon bifurcates, the
larger anterior branch innervating the mantle edge, while
the smaller branch runs posteriorly to the mantle wall in
the region of the anus.
A third large nerve, the left segment of the pleuro-
visceral loop, arises from the right pleural ganglion, runs
diagonally posteriorly dorsal to the esophagus, and enters
the supra-esophageal ganglion on the left side. From this
ganglion arise the left zygoneury, a large osphradiobran-
chial nerve, and the remaining posteriorly directed left
part of the pleuro-visceral loop. The right segment of the
pleuro-visceral loop originates anteriorly from the sub-
esophageal ganglion and runs diagonally posteriorly,
beneath the esophagus.
The pedal ganglia are connected with each other by a
sub-esophageal commissure and with the cerebral and
pleural ganglia on their respective sides by thick connec-
tives. From the anterior face of each pedal ganglion 3
large nerves and one slender nerve arise. The latter, which
has a slightly dorsal origin, and the most lateral of the
thicker nerves innervate the muscles of the basal wall of
the head. The middle of the larger nerves arising from
each pedal ganglion runs directly anteriorly to innervate
Vol. 12; No. 3
Figure 3
The nervous system of Serpulorbis squamigerus
A: lateral view of the cerebral complex
B: dorsal view, pleuro-visceral connectives foreshortened
BG - buccal ganglion
COG - ctenidial-osphradial ganglion;
PIG - pleural ganglion; SbG - sug-esophageal ganglion;
SpG - supra-esophageal ganglion; VGL - left visceral ganglion;
VGR - right visceral ganglion; E - esophagus;
cbc - cerebro-buccal connective; cn - cardiac nerve;
con - ctenidial-osphradial nerves; _ cpc - cerebro-pleural connective;
1- labial nerves; m- mantle nerves; op - optic nerve;
pe - pedal commissure; pd - pedal nerve;
plpc - pleuro-pedal connective; pt- pedal tentacle nerve;
pvcl - left pleuro-visceral connective; pver - right pleuro-visceral
connective; t-tentacular nerve; vis - visceral nerves;
zl- left zygoneury; zr - right zygoneury
CG - cerebral ganglion
PG - pedal ganglion;
ESV EIGER
Page 305
the foot. The medial large nerve on each side innervates
the respective pedal tentacle.
The remaining portion of the nervous system consists
of the posterior segment of the pleuro-visceral loop and its
branches. The right side of this loop runs beneath the
mantle cavity and extends small branches to the mantle
wall and floor. It passes beneath the posterior wall of the
mantle cavity (adjoining the anterior wall of the kidney
sac) and continues posteriorly in close association with
the gonoduct. Below the kidney, where the coelomic gono-
duct turns dorsally to join the pallial gonoduct, the right
pleuro-visceral connective branches, one branch continu-
ing posteriorly, the other running dorsally in the kidney
wall to the right visceral ganglion which lies in the ante-
rior kidney wall directly above the origin of the pallial
gonoduct. From the right visceral ganglion two nerves
arise. One runs to the pericardial region; the other,
forming the posterior portion of the pleuro-visceral loop,
extends to the left to join the left visceral ganglion near
the posterior end of the ctenidium. From the left visceral
ganglion a nerve arises which runs posteriorly along the
ventral surface of the animal into the abdominal region.
A similar nerve, the continuation of the right pleuro-vis-
ceral connective, runs into the abdomen on the right
ventral side of the animal in conjunction with the gono-
duct; it appears to innervate the gonad.
The pattern of the nervous system shown here is typical
for members of the super-family Cerithiacea (see Bouvier,
1887, for numerous other examples). The major variations
here are the retention of unfused visceral ganglia and the
absence of statocysts which occur on the pedal ganglia of
most other gastropods. Considering the sessile habit of
the vermetids, the absence of statocysts is not surprising.
The nervous system of Serpulorbis squamigerus is nearly
identical to that shown for Vermetus triqueter by LacaZzE-
Dutuiers (1860), except that S. squamigerus has a left
visceral ganglion while V. triqueter does not.
BIOLOGY
Serpulorbis squamigerus
MacGinitie & MacGinitie (1968) discussed the habitat,
the gregarious nature, and the mucous-net mode of food
collecting of these snails. Further details from the present
study are added below.
The mucous nets are a product of the extensive paired
pedal mucous glands. These glands are capable of secret-
ing a net as large as 50 cm? in 3 to 4 minutes. The mucus
flows out of the median aperture between the bases of
the pedal tentacles and is carried distally by ciliary cur-
Page 306
THE VELIGER
Vol. 12; No. 3
rents in the grooves on the pedal tentacles. The pedal
tentacles thus spread the single string of mucus issuing
from the aperture of the gland into a triangular sheet
(Plate 45, Figure 1). Some hardening of the mucus ap-
parently occurs on contact with seawater. The density of
the mucus is close to that of seawater since the net is
buoyed upward by very slight turbulence or by only a few
tiny trapped bubbles.
The mucous net remains suspended in the water for 10
to 30 minutes and is then retrieved by the action of the
radula and jaws. The radula is extended far out of the
widely-opened mouth; it grasps the mucous net and pulls
it a short distance down and into the mouth. The jaws
then close on the mucus, holding it during swallowing and
disengagement of the radula. The esophagus of Serpul-
orbis squamigerus is very long and extensible, a modifica-
tion which provides a large storage area for the rapidly
ingested mucous net before its slower passage into the
stomach.
In laboratory aquaria individuals of Serpulorbis squa-
migerus may be stimulated to produce mucous nets by
either strong water currents or agitation of the water. The
addition of dried and ground algae or dried animal mat-
ter (commercial pet fish food) to unagitated water in the
aquaria did not elicit feeding, nor did the addition of live
copepods (Tigriopus californicus BAKER, 1912). It would
thus appear that the usual stimulus for feeding in this
species is water turbulence (the incoming tide?) and not
the presence of food, as BoETTcER (1930) found to be the
case for S. gigas.
Individuals of Serpulorbis squamigerus are predomi-
nantly gregarious, but not exclusively so. They are often
found widely separated from one another on floats in
southern California yacht basins. In these instances the
“communal mess table” emphasized by Morton (1965)
does not occur, but the animals are apparently successful
in their enforced solitary feeding, for isolated individuals
are often of much greater size than aggregated ones.
While the mucous-net method is the predominant meth-
od of feeding in Serpulorbis squamigerus it may not be
the only one; examination of the ciliary currents of the
mantle cavity clearly shows that material filtered out of
seawater by the gills is passed to the oral region. Actual
ingestion of this material was not observed.
Just what constitutes the major food of Serpulorbis in
the field is still uncertain. Laboratory aquaria undoubtedly
present an anomalous situation. However, examination of
nets extended in the aquaria shows them invariably to
contain numerous diatoms, skeletons of small crustaceans,
and unidentifiable debris. In paraffin sections through the
stomach region of S. sqguamigerus taken in the field I have
seen diatoms and occasional small crustaceans and worms.
Whether or not the snails are capable of digesting animal
matter remains open to investigation. The presence of a
crystalline style would appear to argue against such a
possibility (YonceE, 1930, 1932).
MacGiniTigE & MacGinitie (1968) state that Serpul-
orbis squamigerus is preyed upon by the carnivorous pro-
sobranchs Ceratostoma foliatum (GmeEtIn, 1791) and
Shaskyus festivus (Hinps, 1844). In the present study no
predators have been seen, but the presence of trematode
cercariae in the abdomen of S. squamigerus has been
noted frequently. These cercariae appear to feed exclusive-
ly on the gonad of the snail and may cause nearly total
castration. An occasional commensal of S$. squamigerus is
a pinnotherid crab, tentatively identified as Opisthopus
transversus RATHBUN, 1893. These crabs occur in the
mantle cavity, and when removed from one snail have
been observed to re-enter the mantle cavity of another host
quite readily. The snails exhibit little response to the ent-
rance of the crab. Curiously, this association has only been
noted in snails collected from the Malibu region of Cali-
fornia and never from animals collected to the north or
south of this area. The pinnotherids were noted in indi-
viduals of S. squamigerus collected in March and May
1966.
As might be expected, the numerous spaces between the
tubes of large masses of Serpulorbis provide refuge for
great numbers of other invertebrates. Among these are
calyptraeid gastropods, nemerteans, sipunculids, poly-
chaetes, and the ubiquitous nematodes. The tips of the
vermetid tubes are frequently encrusted by bryozoans,
sponges, and compound ascidians.
Explanation of Plate 45
Figure 1: Serpulorbis squamigerus feeding (1)
Figure 2: Petaloconchus montereyensis: mass of living individuals
(X 2.5)
Figure 3: Preserved specimen of Serpulorbis squamigerus seen from
the right side (X 1.5)
Figure 4: Preserved specimens of Serpulorbis squamigerus, male
and female, seen in dorsal view (X 1.5)
C - columellar muscle; F - foot; H - head;
MM - mantle margin; Mn - mucous net;
PS - pallial slit of female; Pt - pedal tentacle;
1, 2, and 3 represent gross body regions: 1, mantle cavity; 2,
region of heart, kidney and stomach; 3, region of digestive gland
and gonad
THE VELIGER, Vol. 12, No. 3 [HapFIELp] Plate 45
Figure 3 Figure 4
Vol. 12; No. 3
THE VELIGER
Page 307
Just how dense the populations of Serpulorbis squami-
gerus may be in some localities was illustrated by PEQUE-
GNAT (1964) who reported concentrations of this species
of 650 per m’. The greatest concentrations of S. squami-
gerus seen in the present study are on a small sandstone
outcropping adjacent to the beach at the Kerkhoff Marine
Laboratory, Newport Bay, California. The stone is sur-
rounded by muddy sand which limits the spread of S.
squamigerus, but nearly every square inch of exposed rock
is covered by them. Sponges and bryozoans are also abun-
dant on and around the vermetid tubes.
KEEN (1961, p. 203) gives the range for Serpulorbis
squamigerus “from Monterey, California, to southern Baja
California.” This is undoubtedly more accurate than the
listing by R. T: Appott (1954), “Forrester Island, Alaska
to Peru,” since many of the reports on which ABBorTtT’s
range is based are by authors who confused S. squamigerus
with other vermetids, some not even of this genus (OLD-
ROYD, 1924, describes “Aletes squamigerus” as a species
bearing an operculum!). Nowhere north of Point Con-
ception, California have large numbers of S. squamigerus
been seen, and on the Monterey Peninsula they are rare.
Within the range given by KEEN (1961), the vertical
distribution of Serpulorbis squamigerus is poorly known.
Studies on the biota around sewer outfalls and off-shore
oil wells (TURNER et al, 1962, 1965a, 1965b) include
records of S. squamigerus from the intertidal zone to
depths of 80 feet. A more reliable maximum depth is prob-
ably 50 feet, since the 80 foot record was of a single
specimen (a shell?) found on a sandy bottom, an unlikely
habitat for a vermetid. In the area studied by PEQUEGNAT
(1964), which consisted of two sandstone reefs offshore
from Corona Del Mar, California, S$. squamigerus was
noted to occur on the reef tops, average depth 11 m, in
areas of greatest turbulence.
Petaloconchus montereyensis
Petaloconchus montereyensis is known only from concho-
logical faunal lists and Morton’s (1965) partial descrip-
tion of the animal based on preserved material. Specimens
used in this study were collected from the type locality,
“Monterey, California” (Dat, 1919), and the surround-
ing coast.
Morton (op. cit.) has suggested that Petaloconchus
montereyensis should be capable of both ciliary filter
feeding and feeding with mucous nets (his judgments
were based on relative size of the ctenidium and pedal
mucous glands). This is indeed the case. The ciliary and
glandular tracts of the gill and mantle walls carry par-
ticles from the gills forward over the mantle floor to the
mesopodial pad in front of the mouth where they are
mixed with mucus and then ingested. However, the pre-
dominant method of feeding is by means of mucous nets
as in Serpulorbis squamigerus. The nets of Petaloconchus
are fragile, and in aggregates of these snails they are fused
in a thin veil which overlies a whole portion of the colony.
The nets are so thin as to be barely discernible unless they
are laden with detritus particles. The nets are produced
and retrieved in a manner precisely like that described
for Serpulorbis.
Petaloconchus montereyensis, while predominantly gre-
garious (Plate 45, Figure 2), is represented occasionally
by isolated individuals. As these individuals have been
found in reproductive condition and equal in size to in-
dividuals occurring in gregarious masses, it appears there
is no great dependence on communal feeding nets.
Petaloconchus montereyensis serves as host to parasites
and commensals. A high rate of infection by trematode
cercariae occurs in populations at Mussel Point, Pacific
Grove, California. The cercariae live on the tissue of the
gonad and may effectively castrate the host. Petaloconchus
is also the host for a pyramidellid snail of the genus Odo-
stomia (Chrysallida). This small parasite has been observed
frequently on masses of Petaloconchus montereyensis tubes.
Odostomia feeds by creeping up to the mouth of a ver-
metid tube and piercing the fleshy edge of the mantle
tissue with its proboscis.
Amongst the masses of Petaloconchus tubes numerous
other small invertebrates find refuge. Phascolosoma agas-
sizit KEFERSTEIN, 1866 is common here; a small clam,
Kellia, occurs frequently, and many other marine snails
are encountered in this habitat. An almost invariable asso-
ciate of P montereyensis is the polychaete Dodecaceria
fistulicola Exuters, 1901. The calcareous tubes of this
worm are generally found in a band above the zone of
Petaloconchus ; no clear-cut demarcation between the pop-
ulations of the two species exists, and often there is a
horizontal band a few centimeters wide where their tubes
are intermixed. Finally, a small ostracod is often found
crawling in and around the tubes of P montereyensis. It
lays its small, lens-shaped egg capsules on the center of
the operculum of Petaloconchus.
Petaloconchus montereyensis may be unique among the
Gastropoda in its habit of periodically producing a new
operculum and moulting the old one. While marine snails
have been found capable of regenerating the operculum
(Hanxo, 1913), I know of no other instance where total
operculum replacement represents a regular activity on
the part of a snail. In Petaloconchus montereyensis gross
appearances suggest that most of the pedal surface has
the capacity to produce opercular material. The opercu-
lum of a fully grown animal is only slightly smaller in
diameter than the tube aperture. It is a flat plate with a
central nucleus and bears a high, raised flange which
spirals outward from the nucleus in a counter-clockwise
Page 308
THE VELIGER
Vol. 12; No. 3
Figure 4
The operculum of Petaloconchus montereyensis
A: the complete operculum
B and C: two successive stages in operculum replacement
Op - operculum
direction (Text figure 4). The fully developed operculum
has a flange which completes three whorls from the nuc-
leus to the outer edge. Marginally the thin outermost
whorl of the flange overlaps the lateral edges of the foot.
Small and intermediate sized individuals of Petalocon-
chus montereyensis are often found in the process of re-
placing a perfectly formed operculum with one only slight-
ly larger. This observation suggests that opercular replace-
ment (rather than simple marginal increment) repre-
sents a natural part of growth in this species. After the
individuals have achieved full growth, the stimulus for
formation of a new operculum may be related to deterior-
ation of the old one. The worn operculum is ragged and
reduced at the edges and along the flanges, and overgrown
by a crust of diatoms and even small red or brown algae.
The process of replacement is first detectable when the
opercular nucleus is displaced to the left on the face of
the foot and the right hand edge of the operculum be-
comes free from the surface of the foot. On lifting the edge
one can observe, lying beneath the old operculum, the
small transparent nucleus of a new one. As the new oper-
culum grows, increasing in size by additions along the
periphery of the spiral, the old operculum becomes pro-
gressively freed from the surface of the foot until its only
attachment is along the extreme left edge. In this stage
the animals often appear to have two functioning oper-
cula. Finally the snail sheds its old operculum by repeat-
edly hooking its edge over the lip of the shell and then
retracting the foot until the operculum is free. New oper-
cula are always recognizable by their smooth glossy sur-
faces, their transparency, and their lack of encrustation
(Text figure 4).
While Petaloconchus montereyensis is known only from
the Monterey Peninsula, California, I have collected
animals from San Juan Island, Washington which are, in
all external aspects, identical to those from Monterey. Its
vertical distribution has never been studied. The species is
most abundant slightly below the zero tide level in areas
of heavy, but broken wave turbulence. Shells of Petalo-
conchus have been collected by Dr. J. H. McLean from
as deep as 90 feet in the Carmel Submarine Canyon, but
I hesitate to assign these to the species P montereyensis on
the basis of the presently available material.
In certain localities (Mussel Point, Pescadero Point, and
Mission Point, Monterey County, California) aggregates
of Petaloconchus montereyensis may completely cover
rock surfaces for areas of two or more square meters. I
have attempted to obtain a rough approximation of the
density of Petaloconchus in such masses by breaking off a
small piece of a large mass, decalcifying the whole piece
and counting all the animals in it regardless of size. The
figure obtained is 12 Petaloconchus per square centimeter,
about half of which are large enough to be sexually
mature. An extrapolation of this figure suggests an aggre-
gation of Petaloconchus covering 2 m’ might easily contain
over 200000 snails.
ACKNOWLEDGMENTS
I am indebted to Dr. Donald P. Abbott for numerous en-
lightening discussions and his editorial helping hand. Dr.
Myra Keen kindly gave of her time and knowledge in
discussions of the taxonomy of the Vermetidae. Mrs. Dana
B. Pearse prepared the line drawings.
This work was supported by PHS grant number 5-F1-
GM-18,937.
Vol. 12; No. 3
LITERATURE CITED
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1954. American seashells. Princeton, New Jersey, D. van
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Bouvier, EucENnE Louis
1887. | Systéme nerveux, morphologie générale et classification
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Dai, WILLIAM HEALEY
1919. | New shells from the northwest coast.
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FRETTER, VERA, & ALASTAIR GRAHAM
1962. British prosobranch molluscs, their functional anatomy
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Hanko, BELA
1913. Uber die Regeneration des Operkulums bei Murex
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Hoimes, SAMUEL JACKSON
1900. The early cleavage and formation of the mesoderm of
Serpulorbis squamigerus CARPENTER. Biol. Bull. 1: 115-121
Keen, A, Myra
1961. A proposed reclassification of the gastropod family
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LacazE-Dutuiers, FELIX Jos—EpH HENRI DE
1860. Mémoire sur l’anatomie et l’embryogénie des vermets
(Vermetus triqueter et V. semisurrectus Putt.) Ann. Sci.
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MacGiniTlE, Grorce Esper & Nettie MacGInITIE
1968. Natural history of marine animals. 2° ed. xii+523
pp.; 286 text figs. McGraw-Hill Book Co., New York
Morton, JoHN Epwarp
1950. Feeding mechanisms in the Vermetidae (Order Meso-
gastropoda) . Nature 165: 923 - 924
1951. ‘The structure and adaptations of the New Zealand Ver-
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Morton, Joun Epwarp
1958. Mollusca. Hutchinson Univ. Libr., London, 232 pp.
1965. Form and function in the evolution of the Vermetidae.
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Oxproyp, Ipa SHEPARD
1924. Marine shells of Puget Sound and vicinity. Publ.
Puget Sound Biol. Sta. 4: 1 - 272
PEguecnaT, WiLus E.
1964. The epifauna of a California siltstone reef.
45: 272 - 283
SAFRIEL, URIEL
1966. Recent vermetid formation on the Mediterranean shore
Ecology
of Israel. Proc. Malacol. Soc. London 37: 27 - 34
SALENSKY, VLADIMIR
1887. Etudes sur le développement du vermet. Arch. Biol.
6: 655 - 759
THIELE, JOHANNES
1931-1935. Handbuch der systematischen Weichtierkunde.
Jena, pp. 1- 1154; 893 text figs.
Turner, Cuarzes H., J.G. Cariiste & Earu E. EBert
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of the Coop. Agreement between the Assoc. and the Calif. Dept.
of Fish and Game: 53 pp. (mimeographed)
Turner, Cuares H., Earu E. Esert & RoBertT R. Given
1965a. Survey of the marine environment offshore of San Elijo
Lagoon, San Diego County. Calif. Fish and Game 51:
81-112
1956b. The marine environment in the vicinity of the Orange
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Ana River Basin Regional Water Pollution Control Board, no.
8, by Calif. Fish and Game Dept. 11 pp. (mimeographed)
YonGE, CHARLES MaurIcE
1930. The crystalline style of the molluscs and a carnivorous
habit cannot normally coexist. Nature, 125: 444 - 445
1932. Notes on feeding and digestion in Pterocera and Ver-
metus, with a discussion on the occurrence of the crystalline
style in the Gastropoda. Sci. Reprt. Grt. Barr. Rf. Exped.
[Brit. Mus. (Nat. Hist.)] 1: 259 - 281
YonceE, Cuarvtes Maurice & E. J. ILEs
1939. On the mantle cavity, pedal gland, and evolution of
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Page 310
THE VELIGER
Vol. 12; No. 3
New Species of Panamic Gastropods
BY
JAMES H. McLEAN
Los Angeles County Museum of Natural History
goo Exposition Boulevard, Los Angeles, California 90007
(Plate 46)
DurING RECENT YEARS Collecting of mollusks in the Pan-
amic Province has greatly increased, due chiefly to the
interest generated by the appearance in 1958 of “Seashells
of Tropical West America” by Dr. Myra Keen. Now that
this book will soon appear in a revised edition, I offer
descriptions of the following new species. Several of the
species are part of the Allan Hancock Foundation collec-
tion now on loan to the Los Angeles County Museum of
Natural History, several have resulted from field work of
the Museum, while others result from the recent collecting _
in the Galapagos Islands by André and Jacqueline DeRoy,
of Isla Santa Cruz, Galapagos Islands.
Repositories of type material for species described here-
in include the following institutions:
AHF — Allan Hancock Foundation (on loan to
LACM)
AMNH_ - American Museum of Natural History,
New York
CAS — California Academy of Sciences,
San Francisco
LACM - Los Angeles County Museum of Natural
History
SDNHM - San Diego Natural History Museum
SU — Stanford University, Stanford, California
USNM_ - United States National Museum,
Washington, D. C.
Arene guttata McLran, spec. nov.
(Plate 46, Figures 1, 2)
Description of Holotype: Shell small for the genus, solid,
umbilicate, turbinate, suture deeply channeled; nuclear
whorls 1}, smooth, rounded, the nuclear tip not raised;
postnuclear whorls 34. Spiral sculpture on the early
whorls consisting of 3 carinations that gradually take
shape on emerging from the undelimited nuclear whorls.
On the body whorl the peripheral carination projects
slightly; this is the 3" of the early spiral cords, above
are 2 strongly beaded cords, between which arises a less
prominent beaded cord, developed only on the final
whorl; below the peripheral cord another strong cord
projects almost as far. The suture is laid above this cord
until the 3" whorl, at which point the suture exposes it;
the base of the cord is not exposed by the suture except
at the upper edge of the mature lip. Base with 6 promi-
nent beaded cords, 2 of which enter the umbilicus. Axial
sculpture of fine lamellae over entire surface, beading
on the spiral cords produced by the thickening and
overlapping of 5 to 8 layers of lamellae in the direction
of growth. Aperture circular, nacreous within, a trace of
denticles within the outer lip. Lip thickened, spiral cords
reflected inward at lip, when seen in basal view. Color
whitish, with random dotting of pink on the spiral cords.
Operculum multispiral, concave, of about 10 whorls,
beaded in a radiating pattern. Height, 4.5 mm, diameter,
5.0 mm.
Type Material: Holotype, LACM 1275; 50 paratypes,
LACM 1276; 3 paratypes, AMNH 154685; 3 paratypes,
CAS 13273; 3 paratypes, SDNHM 51302; 3 paratypes,
SU 9986; 3 paratypes, USNM 679554.
Additional paratypes are available for distribution to
other institutions.
Type Locality: Academy Bay, Santa Cruz Island, Gala-
pagos Islands, Ecuador, 0°45’ S, 90°20’ W, in tidal pool
under rocks. The type lot, consisting of 119 specimens,
was collected by Jacqueline DeRoy, 30 May, 1969.
Referred Material: AHF bottom sample 432, Tagus
Cove, Albemarle (Isabela) Island, 80-100 fathoms, 1
dead specimen, probably of shallow water origin. Al-
though the species is evidently abundant at the type
locality, its absence in collections made at shore stations
by the Hancock Expeditions suggests that its distribution
in the Galapagos Islands is highly localized.
Vol. 12; No. 3
THE VELIGER
Page 311
Discussion: Arene guttata is evidently one of the species
of Arene having a consistent color pattern and on this
distinction alone may be separated from other eastern
Pacific and western Atlantic species known to me. In
addition, other eastern Pacific species having the rounded
periphery are considerably larger and have more numer-
ous spiral cords.
The name is taken from the Latin, guttatus, spotted,
with reference to its spotted color pattern of pink dots.
Arene echinata McLean, spec. nov.
(Plate 46, Figures 3, 4)
Description of Holotype: Shell large for the genus, sturdy
but not massive, rounded-turbinate, umbilicate, suture
deeply channeled. Nucleus of 1 whorl, postnuclear whorls
5, spiral sculpture on the early whorls consisting of 3
strong cords with sharply projecting scales. A 4" strong
spiral cord emerges above the suture on the penultimate
whorl and 2 secondary cords emerge in the channels ad-
jacent to the middle primary cord. On the last whorl
just behind the lip all cords are of nearly the same size
and counting from the suture to umbilicus there are 12
cords, including 2 that spiral deep within the umbilicus.
Axial sculpture of fine growth lamellae, about 7 layers
occur between each 2 projecting scales on the spiral cords.
Aperture circular, nacreous within, outer lip having weak
denticles within. Lip mature but not thickened, abruptly
dipping downward at the suture. Color pinkish brown,
with irregular lighter mottling. Operculum unknown.
Height, 8.5 mm, diameter, 8.0 mm.
Type Material: Holotype, AMNH 154626; 1 paratype,
AMNH 154627; 2 paratypes, LACM 1277.
Type Locality: Espinosa Point, Fernandina Island, Gal-
apagos Islands, Ecuador, 0°16’ S, 91°27’ W. The type lot
of 4 specimens (hermit crab shells) was collected at low
tide by Jacqueline DeRoy, 30 January 1968. Two speci-
mens were originally forwarded to Dr. William K. Emer-
son of the American Museum of Natural History, who
kindly granted me permission to describe the species; 2
additional specimens from the same lot were later received
from Mrs. DeRoy.
Referred Material: AHF bottom sample station 418,
Darwin Bay, Tower Island, Galapagos, 17 fathoms, 1
dead specimen. In addition, dead juvenile specimens are
represented from AHF bottom sample stations in the Gal-
apagos Islands as follows: 438, San Cristobal (Chatham) ;
441, San Salvador (James); and 452, Santa Maria
(Charles), at depths ranging from 24 - 65 fathoms.
Discussion: Avene echinata, another species endemic to
the Galapagos Islands as far as is known, is the largest east-
ern Pacific species of the genus. In this species the usual
beaded sculpture of Arene is spinose to a greater extent
than observed in other species. Mature specimens may
live offshore or may eventually prove to be living at low
tide.
The name is taken from the Latin, echinatus, spiny,
prickly, with reference to the characteristic sculpture.
Vermicularia frisbeyae McLean, spec. nov.
(Plate 46, Figures 5, 6)
Description of Holotype (Turritella-stage): Whorls 12,
overall slope of shell slightly convex on the sides, nuclear
tip lost, first remaining whorl sculptured by 2 strong
raised keels that are equally spaced between the sutures;
by the 9" whorl the posterior of the 2 keels is diminished
in size and prominence; fine spiral threading appears at
about the 10" whorl and the posterior of the 2 keels is
barely perceptible, the lowermost or peripheral keel re-
mains strong at about 4 the distance from the anterior
suture. The base is defined by a strongly projecting keel,
the summit of which is left exposed by the growing edge
of the lip; base nearly flat, sculptured with fine spiral
striae; lip thin (broken on holotype), its growth line trace
slanting backward, forming a broad sinus slightly above
the main carination; inner lip thin, reflected across a
narrow umbilicus. Color whitish with brown maculations
along the lines of growth on the body whorl and base.
Height, 26.8 mm, diameter, 10.8 mm.
Description of Adult Stage: Shell having 3 whorls with
relaxed coil beyond the Turritella-stage; the first of these
whorls in an open spiral, the last 2 with the axis of coiling
at a 45° angle. The basal and the peripheral keels remain
strong and there are 5 less prominent spiral cords on the
outermost side of the whorl; all surfaces bear fine spiral
striae. Height, 61.2 mm, maximum diameter of aperture,
11.0 mm.
Type Material: Holotype, LACM 1278; 1 paratype, LA
CM 1279 (poor condition) ; 1 paratype, USNM 679555;
1 paratype, SU 9988; 2 paratypes, Frisbey collection (1 in
poor condition).
Type Locality: Off the lighthouse, Manzanillo, Colima,
Mexico, 19°03’ N, 104°20’ W, 30 - 40 fathoms. The type
lot, consisting of 6 specimens (Turritella-stage), was
dredged by Mrs. Jeanne Frisbey of Port Isabel, Texas,
February, 1969.
Page 312
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Vol. 12; No. 3
Referred Material: AHF 274-34, Tenacatita Bay, Jalisco,
Mexico, 50 fathoms, 2 specimens ( Turritella-stage) ; AHF
863-38, Bahia Honda, Panama, 30-50 fathoms, 1 specimen
( Turritella-stage); 4 lots, Donald Shasky collection, Red-
lands, California: Gulf of Tehuantepec, Mexico, 45 fath-
oms, 4 specimens ( Turritella-stage) ; La Libertad, El Sal-
vador, 18 fathoms, 1 specimen ( Turritella-stage) ; Gulf
of Fonseca, El Salvador, 38-60 fathoms, 1 specimen (1
free whorl) ; El Salvador, exact locality unknown, 1 spe-
cimen (3 free whorls, described above). Specimens of the
adult stage were examined after the plate had been pre-
pared, too late for illustration in this paper.
Discussion: Vermicularia frisbeyae has a wide distribu-
tion in the Panamic province. It has escaped detection
until now evidently because it lives at depths less frequent-
ly collected by dredging.
The only eastern Pacific or western Atlantic species
of Vermicularia having the large Turritella-stage is V.
fargoi Ousson, 1951, from the Gulf of Mexico, which
occurs at low tide on mud-flats. Vermicularia fargoi is
a variable species but generally has 3 prominent spiral
cords per whorl rather than 2 as in V. frisbeyae.
Turritella willetti McLean, spec. nov.
(Plate 46, Figures 7, 8)
“Turritella sanguinea Reeve, 1849.” -—SwHasxy, 1961, p.
23; pit. 4, fig. 15.
Description of Holotype: Shell moderately large, taper-
ing, whorls 21, with thin periostracal remnants, nuclear
whorls lost; earliest whorls highly convex with deep su-
tures, under magnification the spiral sculpture on about
the 6" whorl consisting of about 16 narrow, raised threads
with slightly broader interspaces; at about the 9" whorl
5 of the threads become more prominent and the area
below the 3"¢ thread from the base bevels in toward the
suture, the cording above tending to become flat sided,
rather than convex as in the early whorls. On the pen-
ultimate whorl 5 strong cords remain, the general effect
is of flat sided whorls, beveling in above and below the
suture. The base is rounded and there are faintly indicated
spiral cords. Aperture nearly circular, lip thin; growth-line
trace slanting backwards, forming a broad, shallow sinus
close to the anterior suture. Color whitish with brown
maculations, stronger cords with alternating light and
dark markings. Height, 73.0 mm, diameter, 13.7 mm.
Type Material: Holotype, LACM 1280; 2 paratypes, LA
CM 1281; 1 paratype, AMNH 154686; 1 paratype, CAS
13274; 1 paratype, SDNHM 51303; 1 paratype, SU 9989,
1 paratype, USNM 679556. Additional paratypes are in
the Shy collection.
Type Locality: Santiago Bay, Manzanillo, Colima, Mexi-
co, 19°06’ N, 104°23’ W, 7-12 fathoms. The type lot was
dredged by Laura and Carl Shy of Westminster, Cali-
fornia, November and December, 1968.
Referred Material: AHF 1087-40, Ensenada de San
Francisco, Sonora, 15-18 fathoms; AHF 947-39, Isabel
Island, Mexico, 15-25 fathoms; LACM A.375, Chamela
Bay, Jalisco, Mexico, 15-40 fathoms; LACM A.375, Tena-
catita Bay, Jalisco, 20-40 fathoms; LACM A.375, Zihu-
Explanation of Plate 46
Figures 1, 2: Arene guttata McLEan, spec. nov. Holotype, LACM
1275. Santa Cruz Island, Galapagos Islands. Height 4.5 mm, dia-
meter 5.0mm x 6
Figures 3, 4: Arene echinata McLean, spec. nov. Holotype, AMNH
154626. Fernandina Island, Galapagos Islands. Height 8.5 mm, dia-
meter 8.0mm x4
Figure 5: Vermicularia frisbeyae McLean, spec. nov. Holotype,
LACM 1278. Manzanillo, Mexico. Height 26.8mm, diameter
10.8 mm X15
Figure 6: Vermiculania frisbeyae, holotype, detail of early whorls
XK 3
Figure 7: Turritella willetti McLEan, spec. nov. Holotype, LACM
1280. Manzanillo, Mexico. Height 73 mm, diameter 13.7mm XX 1
Figure 8: Turritella willetti, holotype, detail of early whorls X 3
Figure 9: Natica brunneolinea McLean, spec. nov. Holotype. LA
CM 1282. Santa Cruz Island, Galapagos Islands. Height 10.8 mm,
diameter 10.5 mm xX 3
Figure 10: Natica brunneolinea. Paratype, LACM 1283. Height
45.8mm, diameter 42.3 mm XI
Figure 11: Natica sigillata McLeEan, spec. nov. Holotype. LACM
1284. Isabella Island, Galapagos Islands. Height 10.0mm, dia-
meter 10.5 mm X3
Figure 12: Natica sigillata. SU 49428. Carmen Island, Mexico.
Height 19.5 mm, diameter 19.2 mm X 2
Figure 13: Cantharus (Gemophos) berryi McLEAN, spec. nov.
Holotype, LACM1286. Banderas Bay, Mexico. Height 21.1 mm,
diameter 11.7 mm XK 1.5
Figure 14: Cantharus (Gemophos) berryi. Paratype, LACM 1286.
Showing egg capsules attached to the shell. Height 20.8 mm, dia-
meter 12.7 mm X 1.5
Figure 15: Fusinus allyni McLean, spec. nov. Holotype, LACM-
AHF 1290. Daphne Minor Island, Galapagos Islands. Height 88.4
mm, diameter 34.2 mm XI
Figure 16: Fusinus allyni. Paratype, LACM-AHF 1291. Height
70.6 mm, diameter 34.6 mm XI
Figure 17: Fusinus allyni. Paratype, LACM-AHF 1291. Height
40.0 mm, diameter 14.3 mm X15
Tue VELIcER, Vol. 12, No. 3
Figure 12
Figure 13
Figure 14
Figure 15
[McLean] Plate 46
Vol. 12; No. 3
atanejo, Guerrero, Mexico, 20-40 fathoms. The last 3 lots
were dredged by the late George Willett in 1938.
Discussion: Turritella willetti is a species familiar to
collectors having dredged along the Mexican coast. SHAS-
Ky (1961) attempted to equate the species with REEVE’s
Turritella sanguinea (REEveE, 1849, p. 6, fig. 27), which
was said to be from “California.” In REEve’s illustration
the final whorls are also convex and other details of
sculpture and color pattern do not correspond. REEVE’s
taxon is most likely not West American.
Turritella willetti, with its strongly convex early whorls,
cannot be confused with other eastern Pacific species, all
of which have fewer spiral cords on the early whorls. The
cording of the mature whorls is variable and is similar
to the pattern observed in T: mariana Dati, 1908. The
largest specimen observed measures 81 mm in length.
Turritella willetti is named for the late George Willett
of the Los Angeles County Museum of Natural History,
first person to collect specimens of the material at hand.
Willett’s collecting in Mexico in 1938 greatly enriched
the Museum’s collections.
Natica brunneolinea McLEan, spec. nov.
(Plate 46, Figures 9, 10)
Description of Holotype: Shell thin, umbilicate, spire
low, whorls rounded, surface of final whorl with thin
yellowish periostracum. Nuclear whorls 3, glossy, first
nuclear whorl dark brown; postnuclear whorls 24, sculp-
tured with fine retractive axial grooves strongest just be-
low the suture and extending only halfway to the peri-
phery. Outer lip thin, umbilicus narrow, partially ob-
structed within by thick spiral ridge terminating on the
inner lip; parietal callus extending slightly forward near
the umbilicus. Color yellowish on upper portion of whorl,
base below the periphery whitish, marked with irregular
brown pencilled lines on the peripheral area only. Operc-
ulum calcareous, white, with 2 deep grooves at the outer
edge. Height, 10.8 mm, diameter, 10.5 mm.
The largest paratype (LACM), which lacks the operc-
ulum, has 4 postnuclear whorls and measures: height,
45.8mm, diameter, 42.3 mm.
Type Material: Holotype, LACM 1282; 1 paratype, LA
CM 1283; 1 paratype, SDNHM 51304; 1 paratype, US
NM 679557; 1 paratype, AMNH 154687.
Type Locality: Academy Bay, Santa Cruz Island, Gala-
pagos Islands, Ecuador, 0°45’ S, 90°20’ W, 50-100 fath-
oms, dredged by André and Jacqueline DeRoy, 26 April
1967 (3 specimens), 27 May 1968 (1 specimen), 9
December 1968 (1 specimen).
THE VELIGER
Page 313
Referred Material: Galapagos Islands, Ecuador: AHF
190-34, S end Albemarle Island, 58-60 fathoms, 3 speci-
mens; AHF 810-38, Barrington Island, 48-73 fathoms, 2
specimens; AHF 814-38, N of Hood Island, 20-40 fath-
oms, 3 specimens; AHF 816-38, N of Hood Island, 50-100
fathoms, 1 specimen with operculum; AHF 817-38, N of
Hood Island, 140-160 fathoms, 1 specimen.
Discussion: Natica brunneolinea reaches a large size; the
paratype measuring 45.8 mm in height is larger than spe-
cimens of other Panamic species examined. It is the only
eastern Pacific species having the color pattern of vertical
brown lines on the upper part of the whorl, hence the
name. The doubly grooved operculum is known in Natica
grayt Puiwippi, 1852, and N. scethra Datu, 1908, but
these species are smaller and have spiral rather than axial
color patterns.
Natica sigillata McLzaun, spec. nov.
(Plate 46, Figures 11, 12)
Description of Holotype: Shell globose, narrowly umbili-
cate, spire low, whorls rounded, surface of final whorl
with thin yellowish periostracum. Nuclear whorls 14,
glossy, yellow brown; postnuclear whorls 34, smooth ex-
cept for fine lines of growth. Outer lip thin, umbilicus
narrow, columellar wall not greatly thickened, spiral um-
bilical cord lacking, a short tongue of white callus extends
slightly across the whorl, but not obstructing the umbili-
cus; parietal callus between this tongue and the upper
edge of the lip not developed, resulting in an uncalloused
band on the parietal wall extending within the aperture.
Color chestnut brown with tent shaped markings of white,
area immediately adjacent to umbilicus white. Operculum
calcareous, white, with 4 raised ridges at the outer edge,
the 2 inner ridges broader than the outer 2. Height, 10.0
mm, diameter, 10.5 mm.
The largest paratype (LACM 1285), which lacks the
operculum, measures: height, 14.9mm, diameter, 13.3
mm.
Type Material: Holotype, LACM 1284; 1 paratype, LA
CM 1285; 1 paratype, SDNHM 51305; 1 paratype, US
NM 679558; 1 paratype, AMNH 154688.
Type Locality: Tagus Cove, Isabella Island, Galapagos
Islands, Ecuador, 0°16’ S, 91°22’ W, 50 fathoms, 4 spe-
cimens, dredged by André and Jacqueline DeRoy, 25 and
29 January 1968. The LACM paratype was dredged at
South Academy Bay, Santa Cruz Island, by the DeRoys,
11 June 1968.
Referred Material: SU 49428, between Carmen Island
and Loreto, Baja California, 20-45 fathoms, Ariel Expedi-
Page 314
tion, 1960, 1 specimen lacking the operculum (Plate
46, Figure 12), height, 19.5 mm, diameter, 19.2 mm.
Discussion: Natica sigillata is the only eastern Pacific
species having a pattern of tent shaped markings. It
differs from most eastern Pacific species in lacking the
axial grooves below the suture and in lacking a thickened
umbilical ridge.
This species must evidently have a wide distribution
although it is presently known only from the Galapagos
Islands and from one specimen from the Gulf of Califor-
nia. The latter specimen shows the large tent markings
only near the suture and on the base, and the general
coloration is yellowish brown rather than dark brown,
but the color may have faded.
The name is a Latin adjective meaning adorned with
little figures or marks, with reference to the color pattern.
Cantharus (Gemophos) berryi McLean, spec. nov.
(Plate 46, Figures 13, 14)
Description of Holotype: Shell of rather small size,
sturdy, spire elevated, whorls rounded, subangulate at the -
periphery. Nucleus of 14 whorls, rounded, brown. Post-
nuclear whorls 6, axial sculpture of approximately 10
low costae per whorl, weakly developed below the suture
and along the canal; spiral sculpture over-riding the axial
sculpture, consisting on the penultimate whorl of 5 nar-
row cords at the crests of broader spiral ridges, with about
5 thin raised spiral lirae between each crest of the spiral
cords. On the final whorl there are approximately 14
cords below the suture, with 6-8 spiral lirae between
each 2 cords. Aperture ovate, canal oblique and broadly
open, aperture and canal extending more than 4 the
length of the shell, outer lip thick, 14 white spiral lirae
within, edge of lip finely crenulate; inner lip well defined
with callus; anal notch slightly constricted, bordered on
parietal wall with a low ridge of callus; siphonal fasciole
ridged around a slight umbilical chink. Periostracum
thin, yellowish, closely adherent. Color yellowish white,
axial ribs dark brown near the crests of the spiral ribs,
axial color missing on the shoulder. Operculum with api-
cal nucleus, dark brown, outer margin yellowish. Height,
21.1 mm, diameter, 11.7 mm.
Type Material: Holotype, LACM 1286; 50 paratypes,
LACM 1287; 25 paratypes, SU 9990; 3 paratypes, AM
NH 154689; 3 paratypes, CAS 13275; 3 paratypes, SD
NHM 51306; 3 paratypes, USNM.
Additional paratypes are available for distribution to
other institutions.
THE VELIGER
Vol. 12; No. 3
Type Locality: Off La Cruz, Banderas Bay, Jalisco,
Mexico, 20°45’ N, 105°30’ W, 10-15 fathoms. The type
lot, consisting of approximately 135 specimens, was
dredged by James H. McLean and Myra Keen aboard the
Sea Quest, 20 to 24 March 1965, guests of Mr. and Mrs.
Richard FE Dwyer, of Corona del Mar, California.
Referred Material: LACM, Banderas Bay, 45 speci-
mens, dredged April 1963, by Cornelius Willis; LACM
65-17, La Cruz, Banderas Bay, 20 specimens collected by
diving on sand bottom, 12 feet depth, James H. McLean,
25 March 1965; LACM A.5498, Chamela Bay, Jalisco,
Capt. Fred E. Lewis, 8 specimens; LACM A.375, Tena-
catita Bay, Jalisco, Mexico, 20-40 fathoms, George apes
lett, 18 February 1938, 9 specimens.
Cantharus berryi is known from a rather restricted area
in the state of Jalisco, Mexico. In recent years it has been
found only along the north side of Banderas Bay, where it
is evidently fairly common on sand and mud bottoms
offshore. I have a specimen on hand reputed to have been
taken by shrimp fishery operations between La Paz and
Mulege on the southeastern side of Baja California, but
the record needs further verification in view of the limited
distribution along the Mexican mainland.
Discussion: Cantharus berryi is the smallest of the east-
ern Pacific species of Cantharus. It is related to C. lautus
(Reeve, 1846) and C. vibex (Broperip, 1833), two other
species having yellowish or whitish shells with darker
coloration along the axial ribs. The egg capsules of this
species are evidently attached directly to the shell (Plate
46, Figure 14). Egg capsules were found on two of the
approximately 200 specimens on hand. ‘To my knowledge,
this has not been observed in other species of Cantharus
(Gemophos). It is characteristic of most if not all species
of Solenosteira Dati, 1890. The eastern Pacific Soleno-
steiras differ in having larger, whitish shells having a
coarse periostracum.
The species is named for Dr. S. Stillman Berry, of
Redlands, California, who has described a number of
species of Solenosteira, among his numerous contributions
to our knowledge of eastern Pacific mollusks.
Fusinus allyni McLean, spec. nov.
(Plate 46, Figures 15 to 17)
Description of Holotype: Shell large, thin, light in weight,
covered with a fine, yellowish periostracum; whorls in-
flated, rounded, except for a sharply projecting periphery,
suture deeply impressed. Apex missing, 9 whorls remain.
Axial sculpture on the 4" whorl of 9, on the final whorl
Vol. 12; No. 3
of 10 low ribs with broader interspaces, crossed on the 4
whorl by 4 and on the penultimate whorl by 6 major
spiral ribs and numerous spiral threads of varying size,
the peripheral spiral cord the most prominent, forming
a keel with rounded projections on the last 3 whorls. Aper-
ture ovate, outer lip thin, simple, but reflecting the spiral
sculpture; columella with a thin white callus. Canal long,
straight, siphonal fasciole lacking, length of aperture and
canal greater than length of the shell. Color whitish, with
reddish brown on the axial ribs between the major spiral
cords. Operculum of holotype withdrawn. Height, 88.4
mm, diameter, 34.2 mm.
A paratype specimen (Plate 46, Figure 16), with
broken apex and canal, but with body whorls of similar
proportions, has a projecting inner lip and a slightly in-
flated outer lip, suggesting that 90 mm is about the maxi-
mum size expected in this species. A paratype specimen
(Plate 46, Figure 17), 40 mm in length, has an intact
apex with 3 smooth nuclear whorls.
Type Material: Holotype, LACM-AHF 1290; 3 para-
types, LACM-AHF 1291; 1 paratype, CAS 13277; 1
paratype, USNM 679560.
Type Locality: Off Daphne Minor Island, Galapagos
Islands, Ecuador, 0°24’30” S, 90°22’40” W, 70-80 fath-
oms, on mud, Velero III station 792 38, 20 January 1938,
6 specimens. Four of the 6 specimens were live-collected,
inci tanty
° all
va
wy)
THE VELIGER
Page 315
2 are subadult and 2 are immature; the smallest of these
is illustrated.
Referred Material: Galapagos Islands: AHF 788-34, SE
of Daphne Major Island, 55 fathoms, 1 broken juvenile;
AHF 201-34, Gardner Bay, Hood Island, 25-35 fathoms,
1 immature specimen 56 mm in length.
Discussion: As far as is known, Fusinus allyni is endemic
to the Galapagos Islands. It differs from all other large
offshore species in the Eastern Pacific in having a more
inflated body whorl but thinner shell and thinner peri-
ostracum.
Fusinus allyni is named for Mr. Allyn G. Smith of the
California Academy of Sciences, in recognition of his
continuing work with the eastern Pacific species of
Fusinus.
LITERATURE CITED
Keen, A. Myra
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi + 624 pp.; illus.
Stanford, Calif. (Stanford Univ. Press)
Reeve, Lovett Aucustus
1849. | Monograph of the genus Turritella.
5: 11 pits.
Suasxy, Donatp R.
1961. Notes on rare and little known Panamic mollusks. The
Veliger 4 (1): 22-24; plt. 4, figs. 11- 16 (1 July 1961)
Conch. Icon.
(May-June 1849)
Page 316
THE VELIGER
Vol. 12; No. 3
Correlation of Radula Tooth Structure and Food Habits
of Three Vermivorous Species of Conus
BY
JAMES NYBAKKEN '
Moss Landing Laboratories, Box 223, Moss Landing, California 95039
and Department of Zoology, University of Washington, Seattle, Washington 98105
(Plate 47)
MEMBERS OF THE GENUS Conus have the most complex
individual radula teeth of any mollusk. Each tooth is
formed of a single sheet of chitin which is rolled as a
sheet of paper is rolled, and is variously ornamented ex-
ternally. The teeth are asymmetrical and difficult to rep-
resent completely in line drawings (Plate 47, Figures 1,
2, 3). For more detailed descriptions of these teeth and
for figures of the variations the papers of Corton (1945),
Prete (1939), WarMKE (1960), and CLENcH & Konpo
(1946) should be consulted. Apparently the rolled chitin
sheet has a considerable amount of plasticity for the
individual teeth of the various Conus species vary greatly
in shape and in type of and arrangement of ornamentation
such that individual species may, in some cases, be dis-
tinguished from congeners on the basis of tooth structure
alone. PIELE (op. cit.) gives an appreciation of the range
of variation found in the genus in his illustrations.
This variation in shape and structure among species
was noted by several workers, but it was not until 1939
that PreLe arranged the teeth into groups of species with
similar structure. Since then many workers have figured
Conus teeth including, in addition to those mentioned
above, BarNnarp (1958), Hanna (1963), ENDEAN & Rup-
KIN (1965), and Von Mot, TurscH & Kempr (1967),
but only ENpEAN & RupkKIN have attempted to relate
a type of tooth structure to a specific prey utilized by the
species. They describe the general shape of the teeth of
fish-eating, mollusk-eating and worm-eating species, using
PIELE’s categories. While they describe in some detail the
structural features common to the teeth of the first two
groups, by far the greatest number of Conus species feed
on polychaetes and other worm-like invertebrates such
* T should like to acknowledge support from NSF grant GB5942X
as enteropneusts (KoHN, 1959; 1968). ENDEAN & Rup-
KIN do not attempt to relate the features of radula
structure of the few vermivorous species they studied to
particular types of prey. They merely note that all vermi-
vorous species seem to have a spur or cone on the base
of the tooth, and that the teeth are rather short and
squat in appearance.
Since the great majority of Conus species are vermi-
vorous and since the teeth of these species have a con-
siderably wider range of interspecific variation in structure
than those in either the known fish-eaters or mollusk-
eaters, it is of interest to ask if in fact it is possible to
correlate certain structural features of teeth or entire
tooth structure with particular types of prey organisms.
During the course of an investigation of the food
habits of Conus species from Indonesian waters and from
West American waters, I dissected 6 specimens of Conus
zonatus Hwass, 1792, 2 C. imperialis LinNAgEus, 1758,
and 3 C’. brunneus Woop, 1828 (Plate 47, Figures 4, 5, 6).
The only remains found in the digestive tracts of C. zon-
atus and C. brunneus were the setae of members of the
polychate family Amphinomidae, a rather unusual family
for Conus to prey upon (Koun, 1959). I found nothing
in the 2 C. imperialis, but KoHN (of. cit.) has shown
them to feed primarily upon amphinomids. Since the
radula teeth of C’. brunneus were known to me to be quite
distinctive, I was stimulated to ask if the radula teeth of
C. zonatus and C. imperialis were also similar.
I subsequently checked the radula teeth of Conus zona-
tus and C. imperialis, and they also proved to be highly
distinctive and very similar to those of C’. brunneus
(Plate 47, Figures 1, 2, 3). It thus seems reasonable to
assume that this particular type of tooth structure is
somehow useful in predation on amphinomids. Since this
Vol. 12; No. 3
THE VELIGER
Page 317
tooth structure is unique among the various Conus tooth
types, I would suggest that other Conus species showing a
similar type of tooth can also be expected to feed on
amphinomids. For example, Von Mot, TurscH & KEMPF
(1967) figure the radula tooth of C. regius GMELIN,
1791 which has a structure similar to those of the 3
species discussed here, and thus I would expect it to be a
predator on amphinomids.
The fact that Conus species prey on amphinomids at
all is quite remarkable. The family Amphinomidae is a
relatively small one distributed primarily circumtropically.
The species are commonly known as “fire worms” (Day,
1967). This name derives from the fact that they have
large, long, hollow setae which break easily upon touch
and release contained poison into the lesions. This causes
severe itching-burning sensations in man (CLELAND &
SoutHcott, 1965). Day (1967) gives a concise discussion
of the family and good illustrations of setae and worms.
Why these few Conus species should choose to specialize
in eating such formidable prey remains unknown as do
the mechanisms which allow the Conus to consume the
amphinomid without damage to itself.
Amphinomids usually live under coral rock or other
rock or in crevices in rocks in shallow water. They are
usually fairly large worms which feed on sponges, ascidi-
ans and hydroids (Day, op. cit.).
Amphinomids are rarely, if ever, taken by other Conus
species. KoHn (1959, 1968) does not report amphinomid
remains from any other Conus species and dissection of
1300 Conus of 39 vermivorous species from Indonesia
turned up only a single instance of amphinomid remains
in a Conus outside of the described amphinomid eaters.
This was in a specimen of Conus eburneus Hwass, 1792.
However, the amphinomid eaters occasionally take other
polychaetes. In a sample of 13 C. imperialis from the
Seychelle Islands, one specimen was found to have eaten
an unidentified species of the polychaete family Eunicidae.
Thus far, the few specimens of C. zonatus and C. brun-
neus which have had food in them had only setae of
amphinomids, but the number of specimens examined re-
mains too small to make meaningful statements about
the amount of the diet composed of amphinomids.
Figures 1 to 3 of Plate 47 show that in all 3 amphinomid
predators each tooth has a small barb just back from the
tip, followed by a very distinctive series of two large
barbs about a third of the distance down the tooth shaft.
Conus brunneus has 3 such barbs, C. imperialis and C.
zonatus have but 2. The third barb of C. brunneus appears
to be an accessory barb of one of the 2 large barbs. One
of the latter protrudes at a slight angle from the long
axis of the tooth and bears a set of denticles on its upper
surface (Plate 47, Figures 1, 2, 3). The presence of these
2 barbs and of the denticles on the ridge of one of the
barbs distinguishes the teeth of these amphinomid pred-
ators.
The remainder of the tooth is unornamented, but the
base of the tooth is large with 1 (Conus brunneus) or 2
(C. imperialis, C. zonatus) protruding prominent bumps
which can be termed spurs.
The teeth of these 3 species are also very thick and
massive when contrasted to most other species of Conus,
and the radula sheaths each contain fewer teeth than are
normally found in other vermivorous Conus species (usu-
ally 4 to 6 immediately usable teeth in that part of the
radula sheath adjacent to the esophagus as opposed to
8 to 15 in other vermivorous species that I have investi-
gated). The teeth are large as indicated by a low shell
length to tooth length ratio. This ratio averaged 25.1 for
13 teeth from 3 specimens of C. brunneus, 20.3 for 10
teeth from 2 C. imperialis, and 30.4 for 10 teeth from
C. zonatus. Most vermivorous Conus species have shell/
tooth ratios of from 30 to 70 indicating small teeth
(Koun, 1963).
It is worth noting that the similarity in radula struc-
ture and food habits is not reflected in shell morphology
(Plate 47, Figures 4, 5, 6). Whereas Conus imperialis and
C. zonatus have similar tapering shells with straight sides,
C. brunneus has a convex outline to the body whorl and
is a shorter and more obese shell. It also lacks the striking
color patterns seen in C’. imperialis and C. zonatus. The
aperture is wide in C. brunneus as contrasted with the
other 2 species, and the interior is not partially obstructed
by an internal ridge as is the case in C. zonatus. These
differences in shell morphology would seem to disallow
the use of any shell characters in predicting food in this
group whereas this does not seem to be the case in the
fish eating Conus species, all of which seem to be charac-
terized by a wide aperture. It should be noted that C.
imperialis and C.. zonatus are considered to be closely re-
lated, and the color patterns are often more similar than
the present figures suggest.
This paper is but the first attempt to correlate specific
types of radula teeth with particular prey organisms
among the vermivorous Conus species. The teeth of the
species described here are very distinctive and have been
shown to correlate with a distinctive prey type. However,
the great majority of radula tooth types found in the
vermivorous Conus species have not as yet been related
to specific prey types and other such correlations should be
looked for among these types. It is hoped that this paper
will stimulate this work.
Page 318
LITERATURE CITED
BaRNaRD, KEPPEL Harcourt
1958. Contributions to the knowledge of South African marine
Mollusca. Part I. Gastropoda : Prosobranchia : Toxoglossa.
Ann. So. African Mus. 44 (4): 73 - 163
CLELAND, JoHN B. & R. V. SoutrHcott
1965. Injuries to man from marine invertebrates in the Aus-
tralian region. Canberra: Nat. Health & Med. Res. Coun-
cil Rept. no. 12: 282 pp.
CLENCH, WILLIAM JAMES & YOSHIO Konpo
1946. The poison cone shell. Occ. papers Moll. 1 (7):
52 - 80
Corton, BERNARD CHARLES
1945. A catalogue of the cone shells (Conidae) in the South
Australian Museum. Rec. South Austral. Mus. 8 (2): 229
to 280; 5 plts.; 1 text fig.
Day, J. H.
1967. A monograph on the Polychaeta of Southern Africa.
Brit. Mus. (Nat. Hist.) Publ. no. 656; 878 pp. London
ENDEAN, RoBERT & CLARE RUDKIN
1965. Further studies of the venom of Conidae. Toxicon
2: 225 - 249
THE VELIGER
Vol. 12; No. 3
Hanna, G Datias
1963. | West American mollusks of the genus Conus; II. Calif.
Acad. Sci. Occ. Papers 35:1- 103; plts 1-11 (28 Jan. 1963)
Kon, ALAN Jacogs
1959. The ecology of Conus in Hawaii. Ecol. Monogr.
29: 47 - 90.
1963. | Venomous marine snails of the genus Conus; in: Ven-
omous and poisonous animals and noxious plants of the Pa-
cific area. Pergamon Press, New York; 456 pp.
1968. Microhabitats, abundance and food of Conus on atoll
reefs in the Maldive and Chagos Islands. Ecology 49 (6) :
1046 - 1062
PIELE, A. J.
1939. Radula notes, VIII. Conus.
London 23 (6): 348 - 355
Von Mot, JEAN-JAcQUES, BERNARD TurRScH & M. KEMPF
1967. | Campagne de la Calypso au large des cétes atlantiques
de l’Amerique du Sud (1961-1962) 16. Mollusques proso-
branches: Les Conidae du Bresil. Res. Scient. Camp. Ca-
Proc. Malacol. Soc.
lypso 8. Ann. Inst. Oceanogr. 45 (2): 233 - 254
WarMKE, GERMAINE L.
1960. Seven Puerto Rico cones: notes and radulae. The
Nautilus 73 (4): 119-124
Explanation of Plate 47
Figure 1: Radula tooth of Conus zonatus
Figure 2: Radula tooth of Conus imperialis
Figure 3: Radula tooth of Conus brunneus (scale = 0.5mm)
Figure 4: Conus zonatus; length 58.3 mm, A. J. Kohn coll. no. 4384
Figure 5: Conus imperialis; length 72.2 mm, A.J.Kohn coll. no. 1728
Figure 6: Conus brunneus; length 46.1 mm, J. Nybakken collection
THE VELIGER, Vol. 12, No. 3 [NyBAKKEN] Plate 47
Vol. 12; No. 3
THE VELIGER
Page 319
Observations on the Reproductive Biology
of the Kellet’s Whelk, Kelletia kelletiu
(Gastropoda : Neptuneidae)
RICHARD J. ROSENTHAL
Westinghouse Ocean Research Laboratory, San Diego, California 92121
(Plates 48 and 49; 4 Text figures)
INTRODUCTION
THe KeELiet’s WHELK, Kelletia kelletu (Forses, 1850)
(Plate 48) has been reported along the eastern Pacific
Ocean from Santa Barbara, California, to San Quentin
Bay, Baja California (Appott, 1954, p. 231). Kelletia
kelletu is commonly found off southern California in kelp
beds and on nearshore reefs. Its vertical distribution is
typically subtidal, extending from 2 to about 70m in
depth.
Kelletia kelletu has a relatively large spindle-shaped
dextrally spiralled shell. It is one of the larger gastropods
found in southern California waters. During this study
the largest individual had a shell length of 148 mm.
Although K. kelleti is one of the more abundant species of
the Neptuneidae found in southern California, it has not
been studied extensively and the literature contains only
brief discussions on the biology of this gastropod (Mac-
Ginitie, 1949; LimpaucH, 1955).
This paper is concerned with the reproductive biology
of Kelletia kelletu. Observations were made during 1968
and 1969 both in the experimental tanks of the U.S. Bu-
reau of Commercial Fisheries at La Jolla, California and
while diving on nearshore reefs off San Diego County.
MATING BEHAVIOR
The sexes are separate in Kelletia kelletu, and mature
males can be identified by the presence of a penis located
in the head region back of the right tentacle within the
mantle cavity. During the reproductive season the mature
testis is usually reddish-brown, and the mature ovary is
yellow-orange. Paired individuals have been sighted as
early as January during 1968 and 1969. Fertilization is
internal and the copulatory act was observed frequently
during March, April and May of both years. During copu-
lation, the male grasps the female’s shell with his foot
and the outer lips of both shells are usually brought close
together. The male then extends his penis over the outer lip
of the female’s shell between the foot and the mantle
cavity (Plate 49).
Mating appears to be somewhat size-selective, since the
female was generally the larger member of the copulating
pair (Text figure 1). The female had a mean shell length
13 mm larger than the male partner in the 124 copulating
pairs observed. The shell lengths in these mating pairs
ranged between 62 - 120 mm for the males and between
71 - 121 mm for the females. Sexually active males greater
than 80 mm in shell length were available to the spawning
population; yet in only 11 out of the 124 copulating pairs
was the male larger than the female. The shell length
(siphonal canal to the apex) was determined by measuring
each individual to the nearest mm with plastic calipers
either underwater or in the laboratory.
Males seem to attain sexual maturity at a smaller size
(earlier age) than do the females. Whether this is the
result of sexual differences in growth rate or due to a
delay in spawning by females until larger size is obtained
has not yet been determined. PEarcE & THORSON (1967)
believe that female Neptunea antigua (LINNAEUS, 1758)
postpone spawning until they have attained full size, and
that males become sexually active at a smaller shell size
than females. Sexual dimorphism in prosobranch mol-
lusks has been observed in various species with the female
generally larger than the male (Hyman, 1967). E>warps
(1968) observed, however, that male Olivella biplicata
(Sowersy, 1825) were significantly larger and grow more
rapidly than the females.
Page 320
120
100
ie}e)
Length of female (mm)
80
70
60 70 80
THE VELIGER
Vol. 12; No. 3
go 100 110 120
Length of male (mm)
Figure 1
A comparison between the lengths of females and males in 124
copulating pairs. These pairs were measured underwater (in depths
of from 18m to 25m) between 11 April and 16 May, 1969.
- = I pair
The size differential in a mating pair may be of benefit
to the female Kelletia kelleti, since she has the ability to
move around more easily with a smaller size male at-
tached to her shell during mating or egg deposition or
both. Distinguishing sexes in the field is usually relatively
easy since the anterior ends of both members are generally
Four locations off San Diego county were investigated: Point Loma
(32°43’N Lat.); Bird Rock (32°49’N Lat.); Point La Jolla
(32°51’30” N Lat.) ; and Del Mar (32°57’ N Lat.).
+ = 2 pairs
pointed in the same direction, with the male on the right
side or on top of the larger female.
Copulation has been observed prior to and during ovi-
position. One female (102 mm) was observed in the labo-
ratory to copulate 6 times with 5 different male partners
over a 30-day period. In each instance, egg laying either
followed or took place simultaneously during copulation.
[RosENTHAL] Plate 48
THE VELIGER, Vol. 12, No. 3
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Vol. 12; No. 3
THE VELIGER
Page 321
SPAWNING BEHAVIOR
Spawning commenced during April both in the labora-
tory and in the subtidal areas examined during 1968
and 1969. The water temperatures at 20 - 25 m for April
1969 in the vicinity of La Jolla, California ranged from
approximately 10.5° C to 13.8° C, compared to 14.5° C
to 16.9° C in the experimental tanks at the U.S. Bureau
of Commercial Fisheries Laboratory.
Orton (1920) pointed out that the temperature gen-
erally controls the process of gonadal ripening in marine
invertebrates, and that to spawn most invertebrates seem
to require much higher temperatures than are necessary
at other periods. The data collected on Kelletia kelletu
suggest that factors other than just temperature are in-
volved in the inducement of spawning. Oviposition in
the laboratory has coincided with spawning in subtidal
regions even though it occurred on the same day at sig-
nificantly different water temperatures — such as 10.5° C
at a depth of 20 m off Point Loma, California, compared
to 16.5° C in the laboratory water table.
Five spawning females which were tagged underwater
(RosENTHAL, 1969) in April 1968 were located in April
or May. of the following year and found to be either
copulating or depositing egg capsules for the second year
in a row. It is believed that spawning takes place annu-
ally in mature females (> 70 mm), and that the greatest
reproductive success is reached when the female is be-
tween 80 - 90 mm in shell length (Text figure 2).
The females deposit their egg capsules on almost any
available hard substrate such as rocks, discarded mollusk
50
eo o -»
a oa
[e}
Number of females
go 100 110 ‘120
Shell length (mm)
Figure 2
The shell lengths of egg laying females. A total of 170 ovipositing
individuals were measured between 11 April and 16 May, 1969
shells, or even on the shells of living Kelletia kelletit.
Oviposition in the aquaria generally was confined to the
vertical sides of the tanks. In subtidal regions the animals
usually favor low relief rocky substrates which are rela-
tively free of sessile macro-organisms. Egg-laying females
have not been observed depositing capsules on substrates
occupied by large numbers of sea urchins.
During the height of the egg-laying period (April to
May) extensive aggregations oi both male and female
Kelletia kelletu have been seen repeatedly in a kelp bed
off Del Mar, California. On 30 April 1968, between 200
and 300 individuals were observed in a spawning aggre-
gation contained within a 20 m? area at a depth of 19 m.
Aggregations of 15 to 20 individuals are quite common
within this kelp bed during the spawning season.
Aggregations of spawning gastropods have been ob-
served in Strombus gigas LinnaEus, 1758 by D’Asaro
(1965), and in Urosalpinx cinerea (Say, 1822) by Car-
RIKER (1955). It has been indicated by Pearce & THor-
son (1967) that adult Neptunea antigua aggregate during
the spawning season. Kelletia helletw is generally found
in an aggregated distribution pattern throughout the year,
but large numbered groups have been observed only dur-
ing communal feeding and spawning activities. It is felt
that mass aggregations of K. kelleti during the spawning
season may be the result of chemoreceptive detection of
the egg capsules or mucous trails of sexually active snails.
Spawned egg capsules may act as additional stimuli
to other mature females which encounter these capsules.
Spawning females seem to favor ovipositing on substrates
which already contain K. kelleti: egg capsules. Spawning
inducement of this sort was observed by D’Asaro (1966)
in the gastropod Thais haemastoma floridana Conrab,
1837. Aside from chemoreception, the egg depositing
stimulus may be influenced by tactile responses in the foot
and head region of mature females encountering egg cap-
sules on the substrate. It seems likely that a series of
factors, such as gonadal ripeness, fluctuating temperatures,
proper substrate, and the presence of other spawning
females may trigger egg deposition in K. kelletit.
In the laboratory a female Kelletia kelletu (104 mm)
was known to spawn on 4 different occasions over a period
of 30 days, depositing a total of 85 egg capsules during
this period. Another spawning female (78 mm) laid 22
capsules over a 24-hour period for a mean per hour rate
of 0.92 capsules. If the female was disturbed in the aquari-
um while egg laying, the spawning act might halt and be
resumed again at some later time. In subtidal areas tagged
female K. kelletit have been observed depositing egg
clusters one day, and on subsequent days these same
individuals were found laying additional egg capsules in
new locations. CARRIKER (1955) observed that spawning
Urosalpinx cinerea females did not always deposit their
Page 322
egg capsules at one time, but may lay a number of egg
clutches throughout the spawning season. Following ovi-
position, the K. kelletii move away from their egg capsules
and there is no attempt to guard or brood these capsules.
EGG CAPSULES,
EGGS, anp DEVELOPMENT
The egg capsule, or ootheca, is the protective enclosure
housing the eggs during the developmental period up to
the time of hatching. The ventral pedal gland in the
middle of the foot manipulates and hardens the capsule
into a definite shape, and secretions are added to cement
each capsule firmly to the substrate by an adhesive gela-
tinous base. Formation and secretion of the stenoglossan
egg capsule has been extensively reviewed by FRETTER
(1941).
The egg capsules are ovoid and somewhat deflated in
shape, one face of the capsule is slightly convex (Plates
48, 49), When first deposited they appear translucent
THE VELIGER
Vol. 12; No. 3
white but darken with the passage of time due to the
development and pigmentation of the larvae within the
egg capsules. The capsules are attached to the substrate by
one end and are usually deposited in groups or clusters,
seemingly without any definite pattern or order. The free
end of the capsule is closed with a proteinaceous plug that
weakens and dissolves as the embryos reach the hatching
stage.
The height of the egg capsule is dependent upon the
size of the spawning female Kelletia kelleti. During labo-
ratory observations a female 78 mm in shell length depos-
ited egg capsules with a mean height of 7.5 mm compared
to a 120mm female which laid capsules 12.3 mm in
height. Individual egg capsules obtained from different
females contained between 400 and 1022 eggs, although
occasionally a few egg capsules in a cluster are empty.
MacGiniTiz (1949) reported counting 2182 eggs in one
K, kelletit capsule.
The newly spawned eggs are yellow in color, each one
between 200 and 300u in diameter. The egg undergoes
a typical spiral determinate cleavage pattern with the
Figure 3
Trochophore stage of Kelletia kelletii after 13 daysin the egg capsule
THE VELIGER, Vol. 12, No. 3 [RosENTHAL] Plate 49
A copulating pair of Kelletia kelletu
E = egg capsules F = female M = male P = penis
5
ih
i
i
EY
P
{
z
=
!
:
/
|
1
a
Vol. 12; No. 3
THE VELIGER
Page 323
Figure 4
Kelletia kelletti veliger larva, 1 day prior to hatching
embryo passing into a trochophore (Text figure 3) and
then a veliger stage (Text figure 4) prior to emergence
from the capsule.
The incubation time seems to be dependent on the
water temperature, although the hatching rate can be
highly variable in different egg capsules developing under
similar water conditions. SCHELTEMA (1967) has demon-
strated the relationship of temperature to the time re-
quired between spawning and emergence of veligers from
their egg capsules in the gastropod Nassarius obsoletus
(Say, 1822). In the laboratory the hatching period for
Kelletia kelletw was 30 to 34 days for capsules developing
in water temperatures between 14.5°C and 17.5° C.
The early embryonic stages are extremely sensitive to
changes in the water temperature — early cleavage stages
were killed when taken from water 16.8° C and placed
into water temperatures between 22.5° C and 24.0° C.
In comparison, egg capsules containing late stage veligers,
when taken from these same water conditions, hatched
after only one day in the elevated water temperatures.
The newly hatched Kelletia kelletii larvae emerge as
free swimming veligers from the opening created by the
dissolving of the gelatinous plug in the free end of the
capsule. From this point they are carried into the water
column to begin a planktonic existence until settlement
is achieved.
ACKNOWLEDGMENTS
I would like to thank the following individuals for assist-
ance with this study: R.E. Bower, J. R. Chess, W.D.
Clarke, R. Lasker, G. Mead and G. Theilacker. I am also
grateful to the U.S. Bureau of Commercial Fisheries
Laboratory, La Jolla, California, for providing laboratory
facilities without which this study could not have been
completed.
LITERATURE CITED
Assott, RoBERT TUCKER
1954. | American seashells. Princeton, New Jersey, D. van
Nostrand Co., Inc.; xiv+541 pp.; 100 text figs.; 40 plts.
CarrikER, MELBOURNE ROMAINE
1955. Critical review of biology and control of oyster drills
Urosalpinx and Eupleura. Special Sci. Rep. U.S. Fish &
Wildl. Serv.: Fisheries no. 148: 1 - 150
D’Asaro, CHartes N.
1965. | Organogenesis, development, and metamorphosis in the
queen conch, Strombus gigas, with notes on the breeding habits.
Bull. Mar. Sci. 15 (2): 359 - 416
1966. The egg capsules, embryogenesis, and early organogen-
esis of a common oyster predator, Thais haemastoma floridana
(Gastropoda: Prosobranchia). Bull. Mar. Sci. 16 (4):
884 - 914
Epwarps, DaLLas CRAIG
1968. Reproduction in Olivella biplicata.
10 (4) : 297 - 304; plt. 44; 3 text figs.
FRETTER, VERA
1941. The genital ducts of some British stenoglossan proso-
branchs. Journ. Mar. Biol. Assoc. U. K. 25: 173 - 211
Hyman, Lippy HENRIETTA
1967. The invertebrates: Mollusca I.
New York; viit+792 pp.; 249 figs.
Koun, ALAN JAcoBs
1961. | Chemoreception in gastropod molluscs.
1: 291 - 308.
LimBAUGH, CONRAD
1955. Fish life in the kelp beds and the effects of kelp har-
vesting. Univ. Calif. Inst. Mar. Res. ref. 55-9; 158 pp.
MacGinirtieE, Georce Eser & Nettie MacGIniriz
1949. Natural history of marine animals.
McGraw-Hill, New York.
The Veliger
(1 April 1968)
McGraw Hill Book Co.,
Am. Zool.
473 pp.; illus.
Page 324
THE VELIGER
Macauass, Hutpa
1948. An ecological study of snails of the genus Busycon at
Beaufort, North Carolina. Ecol. Monogr. 18: 378 - 409
Orton, Joun H.
1920. Sea temperature, breeding and distribution of marine
animals. Journ. Marine Biol. Assoc. U.K. 12: 339 - 366
Pearce, Jack B. « GUNNAR THORSON
1967. The feeding and reproductive biology of the red whelk,
Neptunea antigua (L.) (Gastropoda, Prosobranchia).
Ophelia 4: 277 - 314
ROSENTHAL, RicHarp J.
1969. A method of tagging mollusks underwater. The
Veliger 11 (3) : 288 - 289; 1 text fig. (1 January 1969)
ScHELTEMA, Rupo-r S.
1967. The relationship of temperature to the larval develop-
ment of Nassarius obsoletus (Gastropoda). Bio]. Bull.
132: 253 - 265
TuHorson, GUNNAR
1950. Reproductive and larval ecology of marine bottom in-
vertebrates. Biol. Rev. 25 (1): 1-45.
Vol. 12; No. 3
Vol. 12; No. 3
THE VELIGER
Page 325
The West American Species of Murexiella,
(Gastropoda : Muricidae)
Including Two New Species
EMILY H. VOKES
Department of Geology, Tulane University, New Orleans, Louisiana 70118
(Plate 50)
ON A RECENT COLLECTING TRIP to Panama I was delighted
to be presented with what I thought must be a new species
of Murexiella. To be certain, I checked with Dr. Myra
Keen, Stanford University, the acknowledged expert on
Panamic Muricidae and discovered that this was the spe-
cies she had figured in her Sea Shells of Tropical West
America as “Maxwellia (?) humilis (Broperip, 1833)”
(1958, p. 354, fig. 342). As I had considered a somewhat
different species to be “Murex” humilis Broperip, this led
to further correspondence with Dr. Keen and the ultimate
conclusion that her figured specimen was, in fact, not
“Murex” humilis but the new species described below as
Murexiella keenae.
Although KEEN (op. cit,) referred this shell to the
genus Maxwellia, subsequent work by EMERSON (in prep-
aration) has shown that the type of this group should no
longer be placed in the Muricinae. Both the new species
and “Murex” humilis are more correctly referred to the
genus Murexiella CLENCH & PEREZ FARFANTE, 1945 (type
species: Murex hidalgot Crossr, 1869). This genus is
characterized by having an almost circular aperture with
no anal notch and from 4 to 10 foliaceous varices, which
have the fronds connected by a complex laminated web-
bing.
Study of the new species of Murexiella led to an investi-
gation of the entire West American group of species that
belong to the genus. The group is separable into two
divisions: the Murexiella humilis complex and the Mur-
extella lappa (BropEriP) complex with 7 nominal species
in the first’ and 6 in the second. The members of the
“humilis” complex have a globose body whorl, sharply
constricted into an elongate siphonal canal. The varices
bear elaborate frondose spines. The members of the “lap-
* Another new species described by Emerson « D?Artizio in the
present number of The Veliger brings the total to 8.
pa” group have a biconic outline, with almost no notice-
able break between body whorl and canal. The spines are
short, stubby, and only slightly fimbriated. The species
included in this division are most like Murexiella glypta
(M. SmirH) of the western Atlantic Pliocene to Recent
and all are probably descended from an as-yet unknown
common ancestor. In a recent paper describing a new
species of Murexsul from the Galapagos Islands, EMEr-
son & D’Attitio (1969, p. 325) placed the “lappa” spe-
cies in the genus Murexsul (type species: “Murex”
octogonus Quoy & GarmarD, 1833). Although I agree
completely with these authors that their new species is
a Murexsul, I cannot place the other species in that group.
There is considerable similarity of morphology between
the two, but the intervarical webbing, although greatly
reduced, is still present in the species under discussion.
Also, the coloration and the nature of the aperture with its
small, circular opening and raised peristome indicate that
the West American species are closer to Murexiella than
to Murexsul. As I have said before, and will continue to
say, we must never forget that all supraspecific groups are
completely artificial and the boundaries between the ar-
bitrarily separated “natural groups” must be gradational.
For this reason there will always be those intermediate
species that might be just as well placed in either of two
adjacent groups. The species of the “lappa” complex are
in that position.
“Murex” humilis was originally described from Santa
Elena, Ecuador (Broperip, 1833, p. 175), and was first
figured by G. B. Sowersy, Jr. (1834, plt. 65, figs. 46, 47).
These figures show a shell with a marked brown spiral
band circling the body whorl at the shoulder. The tips of
the spines and the siphonal canal are brown, with a white
band at the base of the body whorl. The specimen figured
here (Plate 50, Figures 1 - 3) is from off Guaymas, Mexi-
co, and has slightly different coloration. In it the brown
Page 326
shoulder band is confined to the area immediately adja-
cent to the suture, not extending as far as the row of
spines at the shoulder. A second narrow band encircles the
body whorl just at the juncture with the siphonal canal.
Thus what is white or brown in the Ecuadorian shell is
reversed in the Mexican one. The tips of the spines are
brown in some Mexican specimens, although not in the
one figured.
The possession of some sort of spiral brown stripes is
almost characteristic of Murexiella. Most of the West
American species have these stripes developed to a greater
or lesser degree. Some specimens of a species may occur
in an all brown variant, but other specimens of the same
species will be banded. The nature of the coloration, while
consistent in its general appearance, is variable in partic-
ulars. Murexiella hidalgoi, the western Atlantic type of
the genus, has only a faint brown stripe, if any, in the
adult stage, but the juvenile specimens often have a strong
brown marking. Murexiella macginty: (M. Smrru, 1938)
the western Atlantic “analog” of M. humilis, also has these
stripes, which are most clearly indicated on the shoulder
of the shell, appearing as brown blotches between the
varices. This tendency toward brown and white stripes is
found in other muricine genera, including Hexaplex s. s.
and Phyllonotus, and suggests an original common ances-
tor for the three groups.
“Murex” norrisii REEvE, 1845 (Conch. Icon.: Murex,
pit. 28, fig. 129), described without locality data and not
discovered since, is very much like Murexiella humilis.
Murexiella norrisii is a more elongate shell than M. hu-
milis, with 3 small spines in the area between the base of
the body whorl and the first large spine on the siphonal
canal. Some specimens of M. humilis show a trace of these
3 small spines so that M. norrisii may prove to be only an
extreme variant of M. humilis. However, until a specimen
is found that more nearly matches the original material
figured by Reeve, M. norrisii will be retained as a valid
species. There are two specimens of this form in the col-
lections of the British Museum (Natural History), which
may or may not be the types, but they are consistent in
their appearance so that the type cannot be dismissed as
just a pathologic specimen of M. humilis.
Another species of Murexiella described from the Gulf
of California, “Murex” taeniatus SowErsy, 1860 (Proc.
Zool. Soc. London, prt. 27, p. 428; plt. 49, fig. 3), may
well also be a synonym of M. humilis. Kren (1958, p. 361,
fig. 362) figured a large specimen of what seems to be
Murexiella vittata as this species, noting the similarity to
M. vittata at the time. Sowersy’s illustration shows a
much more inflated shell than M. vittata, with 3 broad,
dark brown spiral bands. In his original description he
described the species as “fusco bifasciata,” but changed
THE VELIGER
Vol. 12; No. 3
this to “castaneo trifasciata” in the Thesaurus Conchyli-
orum (1879, p. 29). His illustration shows a shell more
akin to M. humilis than to M. vittata, to my eye, but as
the type was not to be located in the British Museum in
1964, this species must remain a question mark for now.
In fact, SowersBy’s illustration looks so much like the
Atlantic species Murexiella macgintyi, I suspect there may
be an error in the locality.
The fourth species of the Murexiella humilis complex
is “Murex” santarosana Datt, 1905 (Nautilus, vol. 19,
p. 14). This species has been placed in the genus Max-
wellia, but “Murex” gemma Sowersy, the type of
_ Maxwellia, seems more closely related to the genus Aspel-
la. The recently described species Aspella angermeyerae
Emerson & D’Artiuio, 1965, from the Galapagos Is-
lands, clearly demonstrates the link between the 2 groups.
However, “Murex” santarosana is more like the other
West Coast species of Murexiella, as I. discussed in a
previous paper (E. H. Voxes, 1964, p. 14) without ap-
preciating the true implications of the resemblance. The
radula of M. santarosana is identical, however, as well as
several points of shell morphology, to that of Maxwellia
gemma and so it would seem that Murexiella santarosana
is another of those intermediate species. As KEEN (1958,
p. 354) noted, Murex fimbriatus A. Apams, 1853, non
Murex fimbriatus Broccut, 1814, is a synonym of Murexi-
ella santarosana.
The fifth species of the Murexiella humilis group is
“Murex” diomedaeus Dat, 1908 (Bull. Harvard Mus.
Comp. Zool., vol. 43, p. 313; plt. 12, figs. 4, 5), described
from 85 fathoms in the Gulf of Panama. Murexiella
diomedaea * is the West American “analog” of Murexiella
hidalgoi. As a photograph of this specimen has never been
published, it seems desirable to include an up-to-date il-
lustration of this lovely species (Plate 50, Figures 6, 7).
In the collections of the American Museum of Natural
History there is a specimen (no. 92435) of M. diomedaea
taken by shrimpers off Cedros Island, Mexico, extending
the range considerably to the north.
The sixth member of the group is the new species from
Panama mentioned above. It is the largest and most
2 The spelling of this name is somewhat dubious. In, the original
description Datu spelled it diomedaeus; however, on the plate ex-
planation he spelled it diomedae and on the labels with the holo-
type it is spelled the same. The name is derived from Diomedea,
the generic name for the Albatross, after the U.S. Fish Commis-
sion Steamer “Albatross.” According to the ICZN Code the origi-
nal spelling is considered to be correct unless a typographical
error or a lapsus calami is obvious. Incorrect latinization is not
considered as an “inadvertent error’ and is not to be emended.
Therefore, whatever DaLu’s intention may have been, it seems
best to retain the name as originally published, changing to a
feminine termination to agree with the generic name Murexiella.
Vol. 12; No. 3
massive of the group. Both it and the seventh, which is
another new species discovered while I was engaged in
this study, have been confused with Murexiella humilis in
collections, but both represent valid forms. They are de-
scribed at the conclusion of this paper.
The members of the Murexiella lappa complex are all
very much alike and considering the inherent variability
of the genus Murexiella it is probable that not all of the
named species are valid. All originally described as Murex,
in chronological order of description they are:
1. Murexiella exigua (Broperip, 1833) [Proc. Zool. Soc.
London, prt. 2, p. 174, described from Salango, “Co-
lombia” (i. e., Ecuador) ; Sowersy, 1834, Conch. IIl.,
plt. 60, fig. 17.] This tiny shell, measuring % inch in
height, is undoubtedly the juvenile of one of the species
of Murexiella found on the West Coast, probably M.
lappa. In the original description it is stated to have 5
varices and be of a “dirty-white” color but nothing
more is known of the form.
2. Murexiella vittata (BropErip, 1833) [Proc. Zool. Soc.
London, prt. 2, p. 176, from Guayaquil, Ecuador;
Sowersy, 1834, Conch. Ill., plt. 60, fig. 19.] This spe-
cies, while morphologically similar to M. lappa, is con-
stant in the development of an almost black spiral stripe
around the middle of the body whorl, which is visible
on the inside of the shell as well as the outside. There are
also other additional stripes at the shoulder and the
base of the body whorl but they are not seen on the
inside of the shell. This stripe, which gives the species
its name (vittata means striped), can be seen even in
very small individuals. I have one specimen 8 mm in
height that has this stripe well developed.
3. Murexiella lappa (Broperip, 1833) [Proc. Zool. Soc.
London, prt. 2, p. 177, from Santa Elena Bay, Ecuador;
Sowersy, 1834, Conch. IIl., plt. 60, fig. 15.] This species
has a broad, diffuse light brown band circling the body
whorl. As is typical of all species of Murexiella the num-
ber of varices is variable, from 5 to 7 in this case, in
general decreasing with larger size.
4. Murexiella dipsacus (BropErip, 1833) [Proc. Zool. Soc.
London, prt. 2, p. 194, also from Santa Elena Bay;
Sowersy, 1834, Conch. Ill., plt. 60, fig. 20. It is well
to note that the word dipsacus, meaning a teasel or
thistle, is a noun and does not become dipsaca to agree
with Murexiella.] This species is very close to M. lappa,
but is a somewhat more elongate shell and was de-
scribed as having 8 varices. It may be that the apparent
elongation is a result of the extra varix and not a valid
distinction. The color is identical with that of M. lappa,
as is the number and arrangement of varical spines.
THE VELIGER
Page 327
I have never seen a specimen of M. dipsacus, nor have
I ever seen a specimen of M. lappa with 8 varices, and
the two species may well be the same.
5. Murexiella radicata (Hinps, 1844) [Proc. Zool. Soc.
London, prt. 11, p. 128, from San Blas, Mexico; REEvE,
1845, Conch. Icon.: Murex, plt. 30, fig. 148.] The
type specimen of this species differs only in having
somewhat shorter spines than is shown in the illustra-
tion given by Sowersy for M. lappa (the type of M.
lappa is no longer to be found), and is actually the
normal condition for the species. It is unquestionably
a synonym of M. lappa.
6. Murexiella minuscula (M. Soir, 1947) [Nautilus,
vol. 61, p. 54; plt. 2, fig. 8, from the Pearl Islands,
Panama.] Named as a subspecies of M. vittata, this
form was stated by SmirH to differ from M. vittata s. s.
by its “much smaller size,” the holotype measuring 18.5
mm in height. The usual size of M. vittata is about 23
to 25 mm. He did not note that the coloration is com-
pletely different, the shell being dark brown over most
of the body whorl, with a white stripe where M. vittata
has a black stripe. He did not compare his species with
M. lappa, but it may well be the same.
There is one other species of Murexiella to be found
in the eastern Pacific. This is Murexiella perita (Hinps,
1844) [Proc. Zool. Soc. London, prt. 11, p. 129; REEvE,
1845, Conch. Icon.: Murex, plt. 29, fig. 139], which is
unlike either of the groups discussed above. It is most
like Murexiella levicula (Dat, 1889), from the Gulf of
Mexico, but lacks the marked median groove seen on
the spiral ribs of that species. There is no other similar
West Coast shell.
One final species, originally named from “Pacosmayo,
Peru,” which has been synonymized with Murexiella dip-
sacus, is “Murex” peruvianus SowErRBY, 1841 [Proc. Zool.
Soc. London, prt. 8, p. 147; Conch. IIl., plt. 195, fig. 103].
REEVE (1845) placed SowerBy’s species in synonymy with
the neozealandic “Murex” octogonus Quoy & GaIMARD,
1833 (type of Murexsul), but this was refuted by SUTER
(1918, p. 401). According to PonpErR (1968, p. 31) Mur-
exsul octogonus is extremely variable, and if this is the
case, then SowErRBy’s species probably is a synonym of
M. octogonus. In any case, SowERBY’s shell certainly came
from New Zealand and not from Peru for I have seen
several New Zealand specimens that match his illustration
exactly, complete with brown topped spiral ribs. If it is
not the same as Murexsul octogonus then another name is
necessary for there is an older Murex peruvianus, of
Lamarck, 1816, a Trophon. Murexsul cuvierensts FINLAY,
1927, is probably the next available name.
Page 328
Murexiella (Murexiella) keenae E. H. Vokes, spec. nov.
(Plate 50, Figures 8 to 10)
Shell large for the group, nature of protoconch unknown
but probably 14 smooth, somewhat bulbous whorls; 6
whorls in adult teleoconch. Axial ornamentation on neanic
whorls of 7 small varices, becoming from 5 to 7 heavy,
ramose varices on body whorl. Spiral ornamentation of
2 strong cords on all whorls except the last, which has 6
and an additional 2 on the siphonal canal. Where spiral
cords cross varices small recurved, fimbriate spines pro-
duced, those on the canal slightly larger than on the
remainder of the shell. Aperture circular, with a project-
ing, almost entire peristome, broken only at the juncture
of the siphonal canal. Outer lip strongly recurved, and
marked by a series of laminae connecting the spines.
Entire shell covered with minute scabrous growth lines
and small spiral threads between the major cords giving
a granulated appearance to the surface. Siphonal canal
elongated, open, recurved at distal end, terminations of
previous canals projecting as a series of spurs along a
diagonal line to one side of the channel. Color ranging
from peach to light brown, with dark brown blotches
between the varices at the shoulder of the shell only; aper-
ture white. In some specimens the tips of the varical spines
a slightly darker color than the intervarical areas. Oper-
culum muricoid with a basal (abapical) nucleus.
Dimensions of holotype: height 34.3 mm; maximum dia-
meter 22.5 mm.
Holotype: Los Angeles County Museum of Natural His-
tory no. 1259.
Type locality: Venado Beach, Canal Zone (Venado Beach
is just at the western boundary of the Canal Zone, ap-
proximately 5 miles from the entrance to the Canal) ; Bay
of Panama.
Discussion: This species is more massive than any other
species of Murexiella, either Pacific or Atlantic. Murexi-
ella humilis may attain the same dimensions but has a
much lighter appearance at the same size. The spines are
shorter and more ramose in the new species than in M.
humilis or M. norristi and the coloration is different, M.
keenae having only a series of brown markings in the
intervarical areas at the shoulder of the shell. These are
easily seen in Plate 50, Figure 9. So far as is known the
range of M. keenae is from the Bay of Panama north to
Mexico. Dr. Keen informs me she has seen specimens
taken by the shrimp fishermen in Mexican waters and in
the collections of the American Museum of Natural His-
THE VELIGER
Vol. 12; No. 3
tory there is a specimen (no. 154654) dredged by the
Puritan in 6 to 10 fathoms off the Tres Marias Islands,
Nayarit.
As was mentioned above, this is the species that was
figured by KEEN (1958, figure 342) as “Maxwellia” hu-
milis. It was also figured by Reeve (1845, plt. 13, fig.
50) as “Murex” humilis and that author noted: “This
species has been but imperfectly described and figured
hitherto for want of a good specimen. It is a species of
very peculiar character, its pyriform growth, the curiously
recurved bi-squamate structure of the varices, and its
orange-red coloring are features of considerable specific
interest.” However, the original illustrations of “Murex”
humilis given by Sowersy (1834, plt. 65, figures 46, 47)
are actually very well done and easily identified when
compared with specimens.
It gives me great pleasure to name this species in honor
of Dr. Myra Keen, who has done so much for the study
of West American Mollusca in general and the writer in
particular. In addition, I would like to express my grati-
tude to Mrs. Robert H. Stewart of Balboa Heights, Canal
Zone, who collected the type specimen, and Mrs. Ben H.
Purdy of San Diego, California, who provided the com-
parative specimens of Murexiella humilis (including the
one figured) for this study. There is another very fine
specimen of M. keenae, from Venado Island, near Venado
Beach, in her collection.
Murexiella (Murexiella) laurae E. H. VoKEs, spec. nov.
(Plate 50, Figures 4, 5)
Shell small for the group; protoconch of 34 somewhat
bulbous whorls, ending at a small varix; 6 whorls in adult
teleoconch. Axial sculpture of 8 small varices on early
whorls, gradually decreasing to 5 on later whorls of all
specimens seen. Spiral ornamentation of 2 strong, square-
topped cords on all whorls except last; on that whorl there
are 5 cords on the body and 2 more on the siphonal canal.
In the adult shells spiral cords completely obsolete in
intervarical areas, visible only on abapertural side of
varices. Where each spiral crosses the varices short, foli-
aceous, recurved spines developed, and one additional
spine between the suture and the shoulder spine, where
no spiral is present. Spines connected by a complex web-
bing formed by multiple laminae giving a fretted ap-
pearance to the abapertural face of the varices. Aperture
almost circular, slightly crenulated into the varical spines;
no anal notch, columellar lip smooth, free-standing. Si-
phonal canal moderately long, open by a narrow slit,
Tue VE.IcER, Vol. 12, No. 3 [E. H. Voxgs] Plate 50
Figure 2
Figure 7
Figure 8 Figure g Figure to
Vol. 12; No. 3
recurved at the distal end, bearing a series of spurs, which
mark the previous terminations of the canal. Color a rich
chestnut brown over entire shell, subsutural area only
slightly darker. Operculum muricoid, with a basal nucleus.
Dimensions of holotype: height 20.5 mm, maximum dia-
meter 13.5 mm.
Holotype: Los Angeles County Museum of Natural His-
tory no. 1260.
Type locality: Punta de Juluapan, Manzanillo, Colima,
Mexico, 17 fathoms.
Discussion: There are 8 specimens in the type lot of this
new species, all collected by Mr. and Mrs. Carl Shy of
Westminster, California. The species most nearly resem-
bles the other new Murexiella described above but may be
distinguished by its smaller size (both holotypes have 6
teleoconch whorls) and by the fact that the intervarical
areas of M. laurae are completely smooth in contrast to
the ribbed intervarical areas of M. keenae. The details
of surface ornamentation are also different in the two
species, M. keenae being covered with a microscopic gran-
ular type of ornamentation while M. laurae is marked
by a very fine pattern of crossed spiral and axial lines of
equal strength, giving a woven cloth-like texture to the
shell surface. Murexiella laurae differs from M. humilis
in having more foliaceous varices, a smaller size, and again
in the smooth intervarical areas.
This new species is named in honor of Laura (Mrs.
Carl) Shy, who so kindly contributed the type specimen.
Five of the paratypes, the largest of which measures 23
mm in height, remain in her collection. One paratype is
in the collection of Mrs. Ben H. Purdy, and one is in my
collection, both through the courtesy of Mrs. Shy.
I would also like to express my appreciation to Dr.
William K. Emerson of the American Museum of Natural
THE VELIGER
Page 329
History who provided additional material for study and
made several useful suggestions for the improvement of
the manuscript.
LITERATURE CITED
BroperiP, WILLIAM JOHN
1833. Characters of new species of Mollusca and Conchifera
collected by Mr. Cuming. Proc. Zool. Soc. London for
1832: 173 - 179 (14 January 1833)
Emerson, WituiaM KeiTH « ANTHONY D?’ATTILIO
1969. A new species of Murexsul (Gastropoda) from the Gala-
pagos Islands. The Veliger 11 (4): 324-325; plt. 50
(1 April 1969)
Kren, A. MyRA
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi + 624 pp.; illus.
Ponper, W. E
1968. | Nomenclatural notes on some New Zealand rachiglos-
san gastropods with descriptions of five new species. Rec.
Domin. Mus. 6 (4): 29 - 47; plts. 1-5
Reeve, Lovet, Aucustus
1845-1846. Conchologia Iconica; or Illustrations of the shells of
molluscous animals. Monograph of the genus Murex. Lon-
don, plts. 1 - 36 (April 1845 to April 1846)
SoweErBy, GEorGE BRETTINGHAM (2%? of name)
1834-1841. Conchological illustrations. Murex. London, plts.
58-67 (30 May to 1 October 1834) ; plts. 187-199 (1 Janu-
ary to February 1841), together with Murex: a catalogue of
Recent species, 9 pp., London
1879. Thesaurus Conchyliorum: Monograph of the genus
Murex. London, 4: plts. 380 - 403
Suter, HENRY
1913. Manual of New Zealand Mollusca. MacKay,
Wellington, New Zealand, 1 - 1129; atlas of plates (1915)
VoxEs, Emity Hoskins
1964. Supraspecific groups in the subfamilies Muricinae and
Tritonaliinae (Gastropoda: Muricidae). Malacologia 2 (1):
1-41; plts. 1-3 (September 1964)
Explanation of Plate 50
Figures 1, 2, 3: Murexiella (Murexiella) humilis (BRopERIP, 1833)
Height 31.5 mm; maximum diameter 22.5 mm. Off Guaymas, Sono-
ra, Mexico
Figures 4,5: Murexiella (Murexiella) laurae E.H. Voxes, spec. nov.
Holotype: Los Angeles County Museum of Natural History no. 1260.
Height 20.5 mm; maximum diameter 13.5 mm. Punta de Juluapan,
Colima, Mexico, 17 fathoms
Figures 6, 7: Murexiella (Murexiella) diomedaea (Dat, 1908)
Holotype: U.S.N.M. no. 123020. Height 29mm; diameter (ex-
cluding spines) 16.0 mm. Gulf of Panama, 85 fathoms
Figures 8, 9, 10: Murextella (Murexiella) keenae E. H. VoxEs, spec.
nov. Holotype: Los Angeles County Museum of Natural History,
no. 1259. Height 34.5mm; maximum diameter 24.0mm. Venado
Beach, Canal Zone, Bay of Panama
Note: In all figures (except nos. 2 and g) the specimens are
whitened to show details of ornamentation
(all figures X 2)
Page 330
THE VELIGER
Vol. 12; No. 3
Two New Epitoniidae from the Galapagos Islands
(Mollusca : Gastropoda )
HELEN DUSHANE
15012 El Soneto Drive, Whittier, California 90605
(Plate 51)
SPECIES OF Amaea are uncommon in the eastern Pacific
Ocean with but three species having been described:
Amaea brunneopicta (Datu, 1908), A. ferminiana (DALL,
1908) and A. tehuanarum DuSHANE & McLean, 1968.
During the past months specimens of a fourth species
of Amaea from the eastern Pacific have been sent to the
author for identification. Comparison of these shells with
photographs of Amaea (Scalina) retifera (DAL, 1889) in
CLENCH & TuRNER, 1950, from the western Atlantic and
with the holotpe (USNM 83733) of Da.u’s taxon sug-
gests analogous affinity with the specimens from the east-
ern Pacific; this taxon is here described.
Amaea (Scalina) deroyae DUSHANE, spec. nov.
(Plate 51, Figure 2)
Description of Holotype: Shell medium in size, thin,
pale brown, nuclear whorls missing in holotype; remain-
ing 13 whorls strongly convex, with a round shoulder;
suture deeply impressed, aperture subcircular; axial sculp-
ture foliaceous with about 31 costae on the body whorl;
costae scalloped, with high areas at the intersections of
the spiral sculpture; 6 to 8 spiral ridges on the body
whorl; this reticulate pattern is found on all whorls;
within the reticulations are minute axial and spiral
threads; basal area set off by a strongly projecting ridge
and characterized by low but well-defined reticulate sculp-
ture; columella short, terminating in a shallow sinus;
outer lip thin, with about 7 crenulations, umbilicus. lack-
ing; operculum missing in holotype. Dimensions: length
22.8 mm, width 7.3 mm (holotype).
Type Material:
Holotype: Los Angeles County Museum of Natural
History; LACM-AHF 147-34, Invertebrate Zoology Type
Collection, catalog number 1236
Paratypes (5): Los Angeles County Museum of Natu-
ral History; LACM-AHF 147-34, Invertebrate Zoology
Type Collection, catalog numbers 1239, 1240, 1241, 1242,
1243
Paratypes (2): The American Museum of Natural His-
tory; AMNH number 152604
Paratypes (2) : The American Museum of Natural His-
tory; AMNH number 152605
Paratype (1): Mrs. Jacqueline DeRoy collection
Type Locality: Tagus Cove, Isla Isabela (Albemarle Is-
land), Galapagos Islands, Ecuador, 0°16’S Lat., 91°22’
W Long., 6 specimens dredged by the Velero III at sta-
tion AHF 147-34, in 30 fathoms, rock and coral bottom, 13
January 1934. Five additional specimens were dredged by
Mrs. Jacqueline DeRoy at the same locality in 35 to 60
fathoms, 27 January 1968.
Additional Localities: Five specimens in the Santa Bar-
bara Museum of Natural History Collection and one in
the DuShane Collection, referable to this taxon, were
trawled off Cabo Haro, Guaymas, Sonora, Mexico, 27°52’
N Lat., 110°52’ W Long., in 45 to 90 fathoms, 2 Sep-
tember 1960. Two additional specimens in the Shasky
Collection were trawled off the west side of Isla Monser-
rate, Gulf of California, Mexico, in 20 to 40 fathoms, 1
September 1960. One worn specimen in the Shasky Col-
lection was dredged off Punta Final, Gulf of California,
Mexico, in 20 fathoms, January 1961. Two specimens in
the American Museum of Natural History Collection were
dredged off Maria Magdalena Island in the Tres Marias
Islands group on the Puritan-American Museum Expedi-
tion, in 28 fathoms, March 1967. A third specimen in the
American Museum of Natural History Collection was
dredged off the southwest side of Isla San Diego, Gulf of
California, 25°11’30” N Lat., 110°42’40” W Long., on
the Puritan-American Museum Expedition, in 25 to 40
fathoms, 30 April 1957. This specimen appears to be
freshly dead.
THE VELIcER, Vol. 12, No. 3 [DuSuane_E] Plate 51
Figure 1 Figure 2
Figure 1
Epitonium (Nitidiscala) hancocki DUSHANE, spec. nov.
Holotype: Hood Island, Galapagos Islands, Ecuador. Length 13 mm
Photograph X 4; courtesy Los Angeles County Museum of Natural History
Figure 2
Amaea (Scalina) deroyae DUSHANE, spec. nov.
Holotype: Tagus Cove, Albemarle Island, Galapagos Islands, Ecuador.
Length 22.8mm. Photograph X 1.2; courtesy Los Angeles County
Museum of Natural History
Vol. 12; No. 3
THE VELIGER
Page 331
Table 1
Shell Measurements of Amaea (Scalina) deroyae spec. nov.
6 q § 8 gf
4: fas rr
g god FS Locality
Zan AAS S)
Los Angeles County 6 22.8 X 7.3 14 Isla Isabela, Galapagos Islands, Ecuador
Museum - Allan 14 x5 13
Hancock Foundation 13 xX 4 13
specimens 8 X3 10
6 X2 10
Sieowe 9
American Museum 2 16.5 X 5.5 16 Isla Isabela, Galapagos Islands, Ecuador
of Natural History 13. X 4.5 14 (152604)
American Museum 2 5 X2 6.5 Maria Magdalena Island, Tres Marias
of Natural History 3.8 X 1.5 8 Islands, Mexico ((67 : 75204)
Puritan-American
Museum Expedition
American Museum 1 14 xX4 13 Isla San Diego, Gulf of California,
of Natural History Mexico (76220)
Puritan-American
Museum Expedition
Santa Barbara 5 20 X75 14 Cabo Haro, Gulf of California,
Museum of Natural 13 xX 4 13 Mexico
History 12 x3 13
2.5 X 1.3 6
2 X08 5
Dr. Donald Shasky 2 10 x3 13 Isla Monserrate, Gulf of California,
Collection 9 X25 10 Mexico
Dr. Donald Shasky 1 45 X 1.5 9 Punta Final, Gulf of California,
Collection Mexico
DuShane Collection 1 19 X65 14 Cabo Haro, Gulf of California,
Mexico
Discussion: Amaea deroyae appears to be an analogue
of the Caribbean species A. retifera (Dai, 1889)
(CLENCH & TURNER, 1950; p. 243; plts. 96, 106, figs. 1
to 4). Dax, 1902; plt. 30, fig. 9, and Datz, 1903; plt.
76, fig. 9 present good drawings of the holotype of A.
retifera. Amaea deroyae has the same sculptural charac-
ters, but its shell is apparently consistently smaller than
that of the Atlantic species. The Atlantic specimens attain
a maximum height of about 30 mm, although the holo-
type is 13mm; whereas the largest Pacific specimen
measures 22 mm.
None of the Pacific shells seen were taken alive.
CLENcH & TuRNER (1950, p. 245) describe the opercu-
lum of the Atlantic species; they give the range for A.
retifera as North Carolina to Florida, the Gulf of Mexico
and south to the Barbados, in from 13 to 120 fathoms,
with 50 fathoms being the median. It is a relatively
abundant species as indicated by the records presented by
CLENCH & TuRNER (I. c.) They compare Amaea reti-
fera with A. decussata (LaMaRcK) from the East Indies,
suggesting the relationship to be very close.
Amaea deroyae is known to range throughout the Gulf
of California, and to occur off the Tres Marias Islands
and the Galapagos Islands.
The name honors Mrs. Jacqueline DeRoy of Isla Santa
Cruz, Galapagos Islands, Ecuador, an avid collector,
who continues to contribute to our knowledge of the
Galapagan fauna.
The other epitoniid is a remarkable new species that is
referable to the subgenus Nitzdiscala.
Page 332
Epitonium (Nitidiscala) hancocki DUSHANE, spec. nov.
(Plate 51, Figure 1)
Description of Holotype: Shell white, glossy, axial costae
21, continuous from whorl to whorl, with a sharp spine at
the shoulder of each whorl; nuclear whorls 3, smooth,
convex, brown, glassy; post nuclear whorls 9; suture
deeply impressed ; umbilicus small, not hidden by parietal
lip; surface area between costae lacking spiral threads,
whorls rounded; basal cord lacking; aperture oval; outer
lip reflected and formed by the last costae; operculum
thin, paucispiral, yellowish brown in color. Dimensions:
length 13.0mm, width 5.2 mm (holotype).
Type Material:
Holotype: Los Angeles County Museum of Natural
History; LACM-AHF 814-38, Invertebrate Zoology Type
Collection, catalog number 1235
Paratype (1): Los Angeles County Museum of Natural
History; LACM-AHF 802-38, Invertebrate Zoology Type
Collection, catalog number 1238
Paratype (1): Mrs. Jacqueline DeRoy Collection
Paratypes (2) : DuShane Collection
Type Locality: North of Hood Island (Isla Espamola),
Galapagos Islands, Ecuador, 1°21’S Lat., 90°40’ W Long.,
one live-taken specimen dredged by the Velero III at sta-
tion AHF 814-38, in 20 to 40 fathoms, shell bottom,
28 January 1938. One additional specimen dredged north-
west of Isla Santa Maria (Charles Island), Galapagos
Islands, Ecuador, 1°09’S Lat., 90°35’ W Long., by the
Velero III at station AHF 802-38 in 250 fathoms, sand
and shell bottom appears to be the same species although
badly broken; 4 whorls remain. One live-taken specimen
was dredged in 55 fathoms off the north end of Isla Santa
Cruz (Indefatigable Island) , Galapagos Islands, Ecuador,
25 November 1967 and two specimens were dredged in
112 fathoms off the north end of the island in 1969 by
Mrs. Jacqueline DeRoy.
Discussion: Epitonium hancocki differs from all: other
epitoniids known from the west coast of the Americas by
having a tabulate outline, many costae and a glassy ex-
terior. From Epitonium (Asperiscala) kelseyi BaKer,
Hanna & Stronc (1930), which it most resembles, it
differs by having no spiral sculpture and by having a
thinner shell. From E. implicatum Dati «& OcHSNER
(1928) (fossil, probably Pliocene) it differs by having
THE VELIGER
Vol. 12; No. 3
more costae, a sharp spine at the shoulder of each whorl
and a small umbilicus.
This Epitonium is named in honor of the late Captain
G. Allan Hancock who, with his oceanographic research
vessel, Velero III, undertook numerous expeditions to the
tropical eastern Pacific, including the Galapagos Islands,
where specimens of this new species were obtained.
ACKNOWLEDGMENTS
I am indebted to Dr. William K. Emerson for critical
evaluation of the manuscript and to the Los Angeles
County Museum of Natural History for photographs. For
the loan of the holotype of Amaea (Scalina) retifera I
am grateful to the United States National Museum.
LITERATURE CITED
Baker, FREDERICK, G DaLLas HANNA & ARCHIBALD M. STRONG
1930. | Some Mollusca of the family Epitoniidae from the Gulf
of California. Proc. Calif. Acad. Sci., ser. 4, 19 (5):
41 - 56; plts. 2, 3 (15 July 1930)
CiencH, WILLIAM JAMES & RuTH Dixon TURNER
1950. The genera Sthenorytis, Cirsotrema, ... in the West-
erm Atlantic. Johnsonia 2 (29): 221-248; plts. 96 - 107
Mus. Comp. Zool., Harvard Univ. (30 September 1950)
Dati, WILLIAM HEALEY
1889. Reports on the results of dredging, .. . in the Gulf of
Mexico (1877-78) and in the Caribbean Sea (1879-80), by
the U. S. Coast Survey Steamer “Blake”, .. . Report on the
Mollusca, pt. 2, Gastropoda and Scaphopoda. _ Bull. Mus.
Comp. Zool. 18 (2): 1 - 492; plts. 1 - 40
1902. _ Illustrations and descriptions of new, unfigured, or im-
perfectly known shells, chiefly American, in the U.S. National
Museum. Proc. U.S. Nat. Mus. 24: 499 - 566; plts. 27 - 40
1903. A preliminary catalogue of the shell-bearing marine mol-
lusks and brachiopods of the southeastern coast of the United
States, with illustrations of many of the species. Reprint, to
which are added 21 plates not in the edition of 1889. Bull.
U.S. N. M. 37, new ed.: 232 pp., 95 plts. (3 April 1903)
Dati, WILLIAM HEALEY & WASHINGTON HENRY OCHSNER
1928. Tertiary and Pleistocene Mollusca from the Galapagos
Islands. Proc. Calif. Acad. Sci. ser. 4, 17 (4) : 89 - 140;
plts. 2-7; 5 text figs. (22 June 1928)
Fraser, C. McLean 5 ;
1943. General account of the scientific work of the Véelero
III in the eastern Pacific, 1931-41. Part III. A ten-year list
of the Velero III collecting stations (charts 1 - 115). Allan
Hancock Pac. Exped. 1 (3): 259-431. Univ. South. Calif.
Press, Los Angeles, Calif.
Vol. 12; No. 3
THE VELIGER
Page 333
An Ecological Study of Valley-Forest Gastropods
in a Mixed Mesophytic Situation in Northern Kentucky '
BRANLEY A. BRANSON
DONALD L. BATCH
Eastern Kentucky University, Richmond, Kentucky 40475
(7 Text figures; 3 Tables)
INTRODUCTION
THE MOLLUSCAN FAUNA of Kentucky is poorly understood
(BickEL, 1967), practically nothing having been written
concerning the ecology of terrestrial species. Aquatic forms
are even more poorly known. The objects of this report,
then, are to present the findings of an ecological study
carried out in a rather typical Cumberland Plateau val-
ley system, attempt a correlation between mollusks present
and Horton’s (1945) valley classification as modified by
KueuHne (1962), record data which will be useful in
determining the changes that occur as habitat disruption
proceeds, and to compare the faunas of disturbed versus
undisturbed areas.
A possible criticism of a portion of the study, i. e., that
each station was only visited one time and hence during
cold periods snails would be in hibernation whereas during
warm months they would be active, is negated by the
randomness and intensiveness of our collecting method.
Furthermore, the authors do not confuse the concept of
habitat and niche.
Historical Review of Studies
on Kentucky Gastropods
As indicated by BickEt (1967), almost all of the knowledge
extant concerning Kentucky mollusks has been gleaned by
workers outside the state. This is especially true in the case
of snails. During the nineteenth century, Thomas Say and
' Supported by Eastern Kentucky University Faculty Grant 5-3-
372-6
Rafinesque contributed periodic lists and descriptions,
mostly from the Ohio River Drainage. A few other wor-
kers, notably Lea, described new species. During this cen-
tury, very little has been accomplished. BickEL (1965,
1966) conducted some brief ecological studies on some
aquatic species. CLENCH (1926, 1962a, 1962b) and
CLencH & TuRNER (1955) have presented significant
findings in aquatic species, as have RosEwaTER (1959)
and Price (1900). The latter paper also contained infor-
mation on terrestrial species. Because of the extensive
studies of GoopricH (1921, 1929, 1934a, 1934b, 1934c,
1937, 1938, 1940, 1941), no fauna is better understood
than the aquatic pleurocerids.
However, in terrestrial species, excluding the assorted
records appearing in PrtsBry’s monograph (1939 - 1948),
only Conkuin (1957), Husricut (1950, 1958a, 1958b,
1960, 1962a, 1962b, 1963a, 1963b, 1963c, 1964a, 1964b)
and KapLan & MINCKLEyY (1960) have made any attempt
to study the diverse fauna of Kentucky. None of these
works bears directly upon ecology.
History and Description of the Area
The deeply dissected, highly dendritic canyon system
chosen for the study lies in the Cumberland Plateau of
Wolfe and Powell Counties, Kentucky (Figure 1). The
valley floors are mostly underlaid by undifferentiated
Mississippian rocks, whereas the dissections themselves
are through much-interdigitated Lee Sandstone under-
laid by Breathitt formation. The latter formation is ex-
posed on the floors, and it consists mostly of gray mica-
careous siltstones, subgraywacke sandstone, dark and light
claystones containing ironstone concretions, some lime-
stone (very sparse), chert, and coal (BRANSON & BatcH,
Page 334
Byartee
sequre 7
sea
roa
Sy
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recat
eas
ay
Figure 1
Map showing the Position of Wolfe and Powell Counties, Kentucky.
1968). The rugged, narrow valley with steep cliffs is
supported by this formation. The overlying Lee Formation
produces even more massive cliffs, 200 to 300 feet high
(flat-faced and vertical), consisting mostly of resistant
sandstone and siltstone, relatively little clay, ironstone,
and limestone being in evidence.
The canyon system heads approximately 4/5 of a mile
northwest of Pine Ridge, Wolfe County (Figure 2). The
three main canyons were sampled, but the numerous side
systems were not. The Tight Hollow system, consisting of
first, second, and third order canyons, drains 0.558 square
mile, and it is three-pronged, roughly Y-shaped. The
south arm, which receives a short, deep bifurcation, heads
at 1040 feet mean sea level (msl), and extends 2.4
miles to its confluence with the north arm. The head-
water cliffs average slightly more than 200 feet in vertical
height. Below this point, the cliffs are of variable height,
THE VELIGER
Vol. 12; No. 3
but always steep, and the enclosed valley is narrow and
rocky.
The northern arm heads at 1240 msl, then extends two
miles to its confluence with the southern or lower one at
1030 feet. From this point, the lower arm extends 2225
feet to make contact with the Mill Creek system at 900
feet msl. Mill Creek meanders between steep walls for
slightly in excess of 12000 feet to open into the much
wider canyon of the Middle Fork of Red River, picking
up Doe Branch, Black John Creek, and Doublecave Branch
en route.
The floor of these canyons is occupied by spring-fed
perennial streams (see “collecting station” notes). Table 1
presents the physical and chemical characteristics of the
streams. In Tight Hollow, the creek is small, ranging from
4 to 6 feet in width and from 1 to 6 inches (excluding a
few pools) in depth at the headwaters. In the second order
stretch, the creek widens to about 15 feet and ranges from
8 inches to 3 feet in depth. Below station 4, the creek runs
in a bed averaging 30 feet in width, but the actual water
course ranges from 30 inches to 15 feet, mostly very shal-
low, clear water. At the point where stations 4 and 5 join,
there is a large pool of approximately 30 feet in diameter,
14 feet in depth.
In the third order canyon, the stream averages 20 feet
in width, 3 to 4 inches in depth, with pools up to 3 feet
deep. Numerous small springs join the stream in this
stretch, and the stream bed drops rapidly, about 12 feet
in 30 linear feet. At the lower end of this station, there
is a pool approximately 40 feet in diameter, and about 44
feet deep, above which is a small waterfall. Below station
6, the stream is about 20 feet wide and varies from 6 to
12 inches in depth.
The portion of Mill Creek occupied by station 8 aver-
ages 12 feet in width and about 2 feet in depth. In the
stretch designated as station 9, the stream has a tendency
to become braided in places, but in general the channel
averages about 17 feet in width and ranges from 3 inches
to 3 feet in depth. Station 10 contains a rapidly dropping
bed, and Mill Creek widens to about 30 feet, alternating
between 2-foot pools and shallow riffles. Most of station
11 is occupied by Mill Lake (Figure 2). Above the lake,
the stream is 25 to 30 feet wide and 14 inches deep, where-
as at the earth-fill dam it is 65 feet deep.
The remaining 3 stations are in the fourth order canyon
of Middle Fork of Red River. Here, the stream ranges
from 30 to 50 feet in width and from 6 inches to 4 feet
in depth.
The stream bottom in Tight Hollow consists of flat
sandstone, deposits of sand, some organic debris, and
sandstone boulders. In the Mill Creek canyon, the bottom
Vol. 12; No. 3 THE VELIGER Page 335
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Figure 2
Map showing the Position of Collecting Stations in Tight Hollow,
Mill Creek, and Red River Canyons, Wolfe and Powell Counties,
Kentucky. Arabic numerals demark stations; Roman numerals indi-
cate canyon-stream order (Horton System). Collections made
midway between arrows (see text).
is primarily flat sandstone, small boulders, and large
amounts of sand. The Red River bottom consists primarily
of sand, gravel, and medium to small rocks, except at
station 14, where large quantities of mud have accumu-
lated.
Some of the general biotic conditions of these canyons
were indicated by BRANSON & BatcH (1968, 1969, and
in press). The upper part of the system has never been
occupied by man, notwithstanding a few abortive at-
tempts to locate silver there at the tum of the century.
Conditions are essentially relict and unmodified from, and
including stations 1 through 5. The dominant trees are
beech, eastern hemlock, tulip tree, red maple, northern
dogwood, and umbrella magnolia on the south ridge;
American holly is abundant from station 6 upgrade. From
station 1 through 5, Rhododendron is exceedingly dense,
_up to 2117 main stalks per acre. Below station 5, the lat-
ter form is progressively replaced by mountain laurel. A
similar relationship exists between hemlock and tulip trees,
and hemlock versus pines (White and Virginia).
These conditions reflect the effect of commercial logging
some 20 years ago. Below station 5, the valley has been
logged, and at the junction of stations 7 and 8 the remains
of an old farm are visible. Ecological conditions on this
point are best described as old field.
Below Mill Lake, which is a trout-stocked fishing lake,
Red River stations 12, 13, and 14, lie in the much-dis-
turbed Natural Bridge State Park. Thousands of fishermen,
campers, hikers and picnickers visit the area annually.
Of the ground vegetation, various species of Polypodi-
aceae, Sedum, Lygodium, Lycopodium, and numerous
mosses are the most important in the undisturbed upper
stations, with some dense stands of Equisetum along the
creek, and thick growths of Mitchella repens LINNAEUS,
1756, on the drier slopes. Downstream, several grasses have
invaded the clearings, and Salix grows luxuriantly along
Page 336
the stream margin. In the undisturbed areas, there is a
thick litter of leaves and needles, in addition to many logs
and limbs in various stages of decomposition. Other than
a little encrusting algae, there is no aquatic vegetation in
the stream until Mill Lake is formed. There, a meagre
growth of Nitella occurs in deep water, and sparse accu-
mulations of Spirogyra elsewhere. Below the lake in Red
River, the bottom is without rooted plants until station 14
is reached. As indicated above, the bottom at this station
has considerable mud, and in backwaters considerable
growths of Typha angustifolia LINNAEUS, 1756, and Pota-
mogeton crispus LINNAEUS, 1756 occur.
METHODS
Field sheets and notes were prepared for each station,
which included general descriptions, field numbers, and
the material included in Table 1. The latter data were
secured by means of tests included in “Standard Methods
for the Examination of Water and Wastewater” (ORLAND,
1965). Turbidity was determined by means of a Hellige
Turbidimeter, and water color was assayed by the plat-
inum-cobalt method (Lamar, 1949).
At each of the collecting stations delineated in Figure 2,
a 12-foot wide swath, located at the center of each station,
was hand-collected from one rim of the valley to the
other. Fine-pointed forceps were utilized to secure small to
THE VELIGER
Vol. 12; No. 3
minute specimens, and in some areas a set of graduated
brass sieves were used to sift litter. Information on plant
associations was obtained by means of random swaths.
COLLECTING STATIONS
Collections were secured from 14 sites. In the text follow-
ing, specimens are referred to proper stations by numbers
in parentheses. All specimens, unless otherwise indicated,
are deposited in the Eastern Kentucky University Museum
of Zoology.
Station 1. 26 February 1966. Valley narrow, V-shaped,
between vertical, concave cliffs which rise over 200
feet; several old talus slides present; all boulders and
rocks are sandstone with some small nodules of iron-
stone. Numerous fallen logs present. Vegetation dense,
including mosses, liverworts, ferns; heavy shade by
Tsuga, Magnolia, Rhododendron, and Ilex. Much
ground seepage. Creek 4 to 6 feet wide, clear, running
over bedrock, rubble, boulders, and sand, 1 to 14 inches
in depth, with a few shallow pools up to 2 feet in depth.
Modest current.
Station 2. 5 March 1966. Valley narrow, V-shaped, the
floor heavily littered with massive boulders, flat sand-
stones, and some topsoil washed in from above; forest
litter deep and humid. Other conditions and vegetation
approximately as at station 1. Creek varies from 6 to 15
Table 1
Physical and Chemical Data for Tight Hollow Creek,
Mill Creek, and Middle Branch of Red River,
Wolfe and Powell Counties, Kentucky
(all temperatures in degrees Centigrade)
Station Number
1 2 3 4 5
Hydrogen Ion conc. (pH) 6.98 7.05 7.08 6.85 7.10
O, (ppm) 10.74 11.46 1085 10.93 11.12
% Saturation of O, 80 88 87 97 91
Free CO, (ppm) 0.85 0.50
Phenolphthalein 0 0 0 0 0
Methyl Orange 1720) 16:83) eli, 111883) 7275
Hydroxide 0 0 0 0 0
Carbonate 0 0 0 0 0
Bicarbonates VLOGS Siilel/eem eS Smee
Hardness (Total: Ca & Mg) 52.97 41.54 2669 36.32 19.57
Air Temperature (°C) 2 a) lk 10.05 10.0
Water Temperature (°C) 1.8 3.7 5 9.0 5.8
Soil Temp. (6” depth, °C) 3.0 5.0 6.0 7.5 6.0
Turbidity (ppm) se — 4:25) 43.3) 7.33
Color (ppm) — — — — 15.5
6 7 8 9 10 11 12 13 14
6.91 7.16 698 7.01 656 7.09 7.51 7.27 7.52
10.78 10.97 872 878 818 9.02 878 9.25 8.37
92 91 90 — aS} IG)
0.37 043 0.75 033 030 047 0.00 020 0.57
0 0 0 0 0 0 1.83 0 0
9.75 25.17 11.75 17.83 24.17 42.83 57.33 65.33 43.0
0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 3.67 0 0
9.75 25.17 11.75 17.83 24.17 42.83 55.50 65.33 43.0
14.93 23.68 18.97 26.49 31.23 46.15 259.42 254.14 158.27
10.75 78 197 — Pils) a) BA YAO) af) 2
7.0 60 16.0 — 19.0 285 27.0 285 285
7.0 60 160 — 11.5 — 17.0 225 —
10) 3H} ee) AO) 0.77 036 O41 1.23 1.53
15.5 100 250 <5.0 <50 <5.0 <50 <50 <5.0
Vol. 12; No. 3
feet in width, with several small cataracts over boulders
(lacking in late summer) and a few pools up to 20 feet
in diameter and 2 feet in depth; bottom of sand, boul-
ders, and flat rock impregnated here and there with
small white nodules; heavily overhung by Rhododend-
ron, and several dead Tsuga logs have fallen into the
stream from the steep sides.
Station 3. 12 March 1966. Cliffs less declivitous; valley
U-shaped, now with a narrow flood plain containing
much organic debris. Vegetation and other conditions
similar to those of previous two stations. Creek, slightly
meandering over bed of sandstone, rubble, small boul-
ders, logs and debris, undercuts the cliffs in places, and
some of the banks have fallen into the stream; alter-
nation of shallow riffles with pools (2 to 3 feet in
depth; one 50 feet in length), and a 4-foot fall is
located near the center of the station. At lower end of
the station, the valley again assumes its narrow char-
acter; stream bed inclines at a steeper angle, and widens
to approximately 30 feet. Two small springs join the
main stream at this point.
Station 4. 19 March 1966. Slope of valley walls less steep
than at station 3. A modest flood plain of gravel, mud,
sand, large and small sandstones, and deep organic
litter, from 50 to 100 feet in width attends the stream.
Stream bed about 30 feet wide, the flowing water from
24 to 5 feet in width, alternating between riffles (some
100 feet long) and pools, flowing over slightly terraced
bedrock; much bank undercutting; a 10-foot fall in
lower 4 of station. Vegetation much the same as at
previous stations.
Station 5. 26 March 1966. Nearly identical with station
4. Rhododendron forms canopy above water. Small
amounts of algae in the stream. At the point where
stations 4 and 5 meet, a pool 30 feet wide, 14 feet
deep, and 150 feet long is produced.
Station 6. 2 April 1966. Valley wider at this point, and
the slope of its walls more gentle; Tsuga partially re-
placed by white pine, and Rhododendron by Kalmia;
forest floor more open with less litter. Stream averages
20 feet in width; bottom of bedrock roughed by numer-
ous terraces, set at right angles to long axis, 2 to 12 in-
ches in height; there is an alternation of sandbottomed
pools and riffles; numerous springs fall into the stream.
Below the middle of the station the stream bed declines
rapidly, about 12 feet in a 30-foot stretch, and the bed-
rock contains many washed-out holes; near the lower
end, the drop is about 17 feet in 100. At the point
denoted by the lower arrow in Figure 2, a nearly
circular pool has been gouged out. It is about 40 feet
in diameter, has a sloping sand bottom, and is 44 feet
THE VELIGER
Page 337
in depth. The pool discharges via a short, straight chan-
nel over a 5-foot fall.
Station 7. April 1966. Similar to station 6, but valley
reverts to the V-shaped configuration with vertical
walls. Much dead wood, logs, and sandstone rubble and
boulders. Considerable ironwood, beech, black alder,
and red maple present; practically no Rhododendron.
In the lower 4 of the station, the stream goes under-
ground for approximately 30 feet before emerging. Near
the end of the station, the stream again submerges, runs
underground for nearly 200 feet, and finally percolates
up through sand and gravel to form a pool 4 feet deep,
55 feet long, and 12 feet wide. At the end of the station
(confluence with Mill Creek), the valley widens ab-
ruptly, and a substantial flood margin is produced. On
the left bank, and also partially adjoining Mill Creek,
there is an old field sere, bearing numerous weedy
plants, sumac, willow, young pine, cedar, Lobelia, and
some other flowering plants.
Station 8. 23 April 1966. Mill Creek Valley. Valley wide,
the slope of its walls moderate, with a flood plain of
200 to 300 yards in width gently inclined towards the
creek; it is covered by grasses, sedges, sumac, various
weedy species, dogwood, and a few maples. There are
some boulders, small stones, rubble and organic silt.
The creek, averaging 12 feet in width, runs in a bed of
bedrock, rubble, sand, and gravel. Near the upper
end of the stream, there are several oxbows, 2 to 5
feet in depth, with mud bottoms and weedy banks with
numerous willows.
Station 9. 14 May 1966. Valley continues to widen, its
walls to become lower. Evidence of an old silver mine
on the left, about 100 yards downstream from the sta-
tion’s upper end. Flood plain similar to that of station
8. Stream about 7 feet wide, 3 or 4 inches deep, flowing
over bedrock, gravel, sand, and small boulders. About
200 yards downstream, a large spring wells into the
channel from the south bank. Below this, there are
some large sandstone deposits along the south bank,
and the channel tends to become braided.
Station 10. 22 May 1966. Essentially similar to station 9,
but valley wider. In places, there are deposits of dead
leaves and soil beneath 50-foot cliffs. Stream 20-foot
wide, 3 to 18 inches deep, running over bedrock, sand,
and gravel, alternating between pools and riffles at
lower end.
Station 11. 25 June 1966. Valley impounded, mostly
filled by Mill Lake. Around the margins of the lake
are many dead trees, standing and fallen, and the early
stages of succession are visible. Otherwise, the condi-
tions are similar to those at station 9, although the
Page 338
valley is somewhat deeper and narrower. Above the
lake, the stream bottom is of sand, and the water is
about 25 to 30 feet in depth. At the point where the
stream widens out into the lake, the bottom possesses
much dead organic matter, and methane gas continu-
ously bubbles up from the bottom. The lake is 65 feet
deep at the dam. Living trees around the dam are
mostly tulip trees, sycamores, sumac, and hawthorne.
Station 12. 11 June 1966. Middle Fork of Red River
Valley. Valley much wider and, in places, the flood
plain is nearly a mile wide. Agriculture practiced
(mostly corn and tobacco), with numerous examples
of old field succession: grasses, sedges, mullein, and
weedy species. Dominant trees are willow, sycamore,
tulip, hawthorne, some White and Virginia pine, and
a few young Tsuga. The area is much disturbed. The
river varies from 20 to 30 feet in width, with pools 3
feet in depth, and long stretches of six-inch water
running over gravel, sand, small rocks, and beer cans.
Station 13. 2 July 1966. Valley approximately 1 mile in
width, with a flood plain of approximately 80 feet on
each side of the stream. General vegetation like that at
station 12, but with large accumulations of dead leaves
and drift-deposited twigs and limbs. Stream 20 to 40
feet in width, 2 to 7 feet deep; side pools contain mud,
dead leaves. and organic ooze. The station ends in a
sand-bottomed pool about 3 feet in depth, 45 in width,
and 200 feet in length.
Station 14. 9 July 1966. Valley conditions similar to those
at station 13, but trees more numerous: black walnut,
willow, white pine, ash, box elder, dogwood, redbud,
hawthorn. River banks heavily overgrown by Kudzu.
Stream 30 to 50 feet wide, 24 to 4 feet deep, running
over sand, mud, and gravel. A low-water dam is placed
at the upper end of the station (Figure 2), below which
is a pool about 50 feet in diameter; its bottom is
mostly muck, and along the sides are good growths
of Potamogeton, Carex, Spirogyra, and narrow-leafed
cattail.
Accounts of Species
A total of 1703 specimens was secured from the 14 sites
delineated above. Represented in the collections were 12
families, 26 genera, and 52 species; two species are ap-
parently new. These are described (not named) below.
In the list which follows, the first numeral presented is
the station designator; the numbers in parentheses repre-
sent the number of specimens collected at each site.
THE VELIGER
Vol. 12; No. 3
PoLyYGYRIDAE
Members of this family, represented by 3 genera and 14
species, accounted for 47.26% of the total collections,
and it is thus the dominant molluscan taxon in the area,
being approached only by the Zonitidae. Of the total
collection, Stenotrema comprised 15.03%, Mesodon 23.67
%, and Triodopsis 8.57%. These data are broken down
further in Table 2, and additional analysis is presented in
the discussion. This pattern is adhered to throughout the
paper.
Table 2
Percentage-Composition by Polygyrid Species
in a Northern Kentucky Faunule
% of total
Species fauna
Stenotrema evardsi (BLAND, 1856) 4.34
Stenotrema stenotrema (PFEIFFER, 1842) 5.46
Stenotrema angellum Husricut, 1958 0.05
Stenotrema barbatum (Cxapp, 1904) 3.75
Stenotrema leat (BINNEY, 1840) 1.17
Mesodon thyroidus (Say, 1816) 1.52
Mesodon zaletus (BINNEY, 1837) 0.05
Mesodon appressus (Say, 1821) 5.81
Mesodon sayanus (Pirssry, 1906) — 5.51
Mesodon ruglei (SHUTTLEWoRTH, 1852) 10.74
Triodopsts tridentata (Say, 1816) 3.69
Triodopsis fraudulenta (Pitssry, 1894) 1.93
Triodopsis denotata (Férrusac, 1821) 0.82
Triodopsis albilabris (Say, 1816) 2.11
Stenotrema evardsi (BLAND, 1856)
Collecting sites: 1(1), 2(3), 3(1), 4(2), 5(24), 6(6), 7(5), 8(6),
9(15), 10(6), 12(1), 14(4)
This Appalachian snail characteristically occupies habi-
tats along stream margins, valley slopes and mountains,
mostly at elevations greater than 750 feet (ARCHER, 1948).
Most of our specimens were found under decaying wood
and in leaf litter. The species was more abundant in the
middle and lower valleys than in the narrow portions
above station 5 (0.9% to 2.56% from station I through
station 4; 4.81% to 15.38% from station 5 through station
11). Below station 11 there was a sharp drop in numbers
(1.78% to 3.33%). These latter stations have been grossly
disturbed, so this may not be a valid observation. On the
other hand, the altitude at the lower end of the study
area is approaching the lower limits of Stenotrema evard-
s?s altitudinal distribution.
Vol. 12; No. 3
Stenotrema stenotrema (PFEIFFER, 1842)
Collecting sites: 1(3), 2(6), 4(7), 5(7), 6(5), 7(10), 8(3), 9(27),
10(5), 11(7), 12(2), 13(2), 14(9)
Stenotrema stenotrema is a rather plastic species as
regards habitat. We secured it in all sorts of places —
beneath decaying hardwood and softwood logs, under
bark, in leaf litter, beneath rocks — and at all altitudes.
In the upper (excluding station 2) and middle stations,
the population varied between 2.91% and 6.79%, reaching
a higher percentage at and below station 9, 7.29% to
9.63%.
Stenotrema angellum Husricut, 1958
Collecting site: 12(1)
The single specimen was found beneath a pile of dead
bark near a tulip tree on a bluff above the stream. This
is essentially the same type of habitat from which Hus-
RICHT (1958) removed the holotype. He (loc. cit.) indi-
cated that the species was often found with S. stenotrema.
However, in collections from other localities, we have
found S. angellum only on the lower slopes, whereas S.
stenotrema is often found at higher elevations as well.
Stenotrema barbatum (Ciapp, 1904)
Collecting sites: 1(1), 2(1), 3(5), 4(2), 5(3), 6(11), 7(24), 9(7),
10(3), 11(6), 12(1)
This species was entirely restricted to the lower sides
of the slopes, where it was found under leaf litter, decaying
wood, and to a less extent, under loose talus. It is difficult
to compare our findings in this area with those secured
elsewhere because of taxonomic confusion between this
form and Stenotrema hirsutum (Say, 1817). This latter
is a smaller species, usually between 6.3 and 6.8mm in
diameter, lacking a buttress on the shell. All our speci-
mens are larger than 6.9 mm (6.9 - 9.0 mm) in diameter,
and all possess a buttress.
From station | through station 5, except at station 3,
Stenotrema barbatum comprised only 9.97% to 1.92%
of the molluscan fauna. At station 5, where the valley
immediately re-assumes its narrow character, the popu-
lation percentage drops to 1.92%. At the middle stations,
6 through 11, the percentage composition ranged from
4.68% to 9.64% (average 7.49% ). In the lower 3 stations,
more or less disturbed, the species represented less than
2% of the total population, or was replaced by the next
species.
Stenotrema leai (BINNEY, 1840)
Collecting sites: 4(3), 6(1), 7(8), 9(6), 11(2), 13(1), 14(3)
Essentially a species of lowland hardwood forests, al-
though sometimes found in coniferous situations, this
species has a rather high moisture requirement and is
THE VELIGER
Page 339
thus limited to stream margins up to around 800 feet
elevation. Bascu, BaINER & WitHM (1961) and others
have found the snail in similar situations. The lowlands
affinities of Stenotrema leai are well-illustrated by these
findings. With the exception of station 4, where a small-
pocket population exists, the snail is completely lacking
in the upper stations (elevations above 1000 feet). From
stations 7 through 14, S. leai comprised from 2% to 3%
of the total terrestrial gastropod population.
Mesodon thyroidus (Say, 1816)
Collecting sites: 9(7), 11(16), 13(7), 14(6)
Pitssry (1940) indicated that Mesodon thyroidus was
a lowland species, usually occupying humid habitats below
650 to 700 feet (occasionally up to 900 feet), and the ob-
servations presented here bear out the premise. Our spe-
cimens were either removed from leaf litter or found
beneath decaying logs. In the lower valleys, M. thyroidus
makes up between 3.5% and 6.25% of the fauna.
Mesodon zaletus (BINNEY, 1837)
Collecting site: 14(1)
Although station 14 is a lowland site, this probably does
not reflect an altitudinal preference. Prtspry (1940) found
Mesodon zaletus up to 2000 feet above sea level in the
southern Appalachians. Since the Lee Formation is very
deficient in calcium, the absence of M. zaletus in most of
this region probably reflects the lack of shell-building
materials, The species is more or less abundant in regions
to the south and west of the Upper Red River Drainage,
and in the lower part of the drainage itself.
Mesodon appressus (Say, 1821)
Collecting sites: 1(2), 2(7), 3(5), 4(12), 5(6), 6(10), 7(9), 8(3),
9(21), 10(5), 11(11), 12(2), 14(6)
According to Pitssry (1940), who quoted a personal
communication from Archer, Mesodon appressus is never
abundant on non-calcareous soils; however, in the upper
valleys, this species was more abundant percentage-wise
than its 2 closest contenders, M. ruglei and M. sayanus,
comprising on the average 8.55% of the fauna (1.94%
to 12.82%). Below and including station 7, the popula-
tion seldom exceeds 5% of the total, and when it does
(11.45% at station 11, for example), the population is
nearly always exceeded by M. ruglei or M. sayanus (see
below). Most specimens were found burrowing beneath
dead leaves or under stones and logs.
Mesodon sayanus (Pitssry, 1906)
Collecting sites: 1(2), 2(1), 4(3), 5(8), 6(4), 7(17), 8(1), 9(9),
10(15), 11(9), 12(1), 13(15), 14(9)
This species develops its largest populations, compared
Page 340
with competitors, in the middle to lower slopes, down to
around 100 feet. From the upper stations to the lower,
there is a steady increase percentage-wise, ranging from
1.63% to 6.20% from stations 1 through 9. Below station
9, except for station 12 (which has been nearly denuded
of trees by agriculture), the percentage composition ranges
from 7.50% through 23.45%. At stations where the per-
centage ran low, the populations of Mesodon ruglet were
larger. Mesodon sayanus is distinctly a forest-floor species.
All of our specimens were found near the bases of hard-
wood trees, along the edges of logs and large boulders
where considerable quantities of leaves had accumulated.
Mesodon ruglei (SHUTTLEWoRTH, 1852)
Collecting sites: 1(5), 2(8), 3(4), 4(3), 5(10), 6(6), 7(53), 8(4),
9(42), 10(1), 11(18), 12(5), 13(10), 14(14)
Mesodon ruglei is the dominant polygyrid in the study
area, and, indeed, one of the most abundant mollusks
present. It seems to have no definite altitudinal preference.
Pitspry (1940) reported it to ascend up to 5000 feet in
other regions. Only at 3 stations (1 (4.85%), 4 (2.91%),
and 10 (1.56%), did its population fall below 5% of the
total, averaging 11.83% (5.26% to 19.87%). The species
likewise was found in various microhabitats: beneath large
and small stones, in dry talus, burrowing in leaves, under
decaying logs, and at the base of clumped grasses on the
flood plains.
Triodopsis tridentata (Say, 1816)
Collecting sites: 1(7), 2(5), 3(6), 4(3), 5(7), 6(2), 7(19), 9(9),
10(1), 11(4)
This Appalachian species was said by Pirssry (1940) to
prefer limestone soils in hilly, shaded terrain and to
avoid plains. However, ARCHER (1942) found Triodopsis
tridentata abundant in sandstone uplands of low relief
with pinewoods cover. These latter findings are essentially
similar to ours, with the exception of differences in
sylvan conditions. From our findings it is apparent that T.-
tridentata produces the largest populations in the uplands
(6.79%, 8.77%, and 15.38% at stations 1, 2, and 3, re-
spectively), becoming much less abundant in the lower
ends of the valleys, except in isolated situations at middle
altitudes (station 7: 6.93%), 1.53% to 4.16% of the
total population. Below station 11 it is replaced by the
following species.
Triodopsis fraudulenta (Pitssry, 1894)
Collecting sites: 12(3), 13(15), 14(15)
Very little is known concerning the habitat requirements
and ecology of this species. It is the dominant Triodopsis
in the lower valleys (5.45%, 13.39%, and 12.50% at the
3 stations), where we found it primarily on afternoon-
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Vol. 12; No. 3
shaded slopes under leaves and decaying wood. KAHN &
Kemp (1930) reported the species as more common on
lower slopes than in the highlands.
Triodopsis denotata (Férussac, 1821)
Collecting sites: 7(4), 9(8), 10(1), 13(1)
This is is another distinctly lowlands species, as shown
by our records. In fact, it was found only on the flood
plains beneath water-deposited debris and logs. At the 4
stations listed above, it was never abundant (1.45%,
3.70%, 1.56%, and 0.89%, respectively), and these find-
ings coincide with the senior author’s observations at
other Kentucky localities. As shown in Table 2, Triodopsis
denotata comprised only 0.82% of the total molluscan
fauna, only slightly more abundant than Stenotrema an-
gellum and Mesodon zaletus.
Triodopsis albolabris (Say, 1816)
Collecting sites: 1(1), 4(9), 5(2), 6(3), 7(8), 8(2), 9(5), 10(2),
11(3), 12(1)
With the exception of station 4, where the population
amounted to 8.73% of the total, Triodopsis albolabris is
not abundant at any sampled locality in the valley system
(0.97% to 3.27%). Pitspry (1940) found the species
to be scarce on sandstone substrate in the Catskills. Our
specimens were found beneath accumulations of leaves
in rock crevices, at the bases of trees, and beneath logs.
HAPLOTREMATIDAE
Haplotrema concavum (Say, 1821)
Collecting sites: 2(1), 4(1), 6(3), 7(20), 8(7), 9(11), 10(1),
11(9), 12(10), 13(7), 14(11)
This is a common species of bluffs overlooking streams,
especially on the lower slopes where leaf mold and woody
debris abound. It is heavily preyed upon by shrews and
Peromyscus, and in turn preys upon polygyrids. The pop-
ulations were relatively small in upland situations, ranging
from 0.97% to 2.63% from station 1 through station 7.
From station 8 downward, the populations were much
larger, sometimes attaining dominant numbers, such as
those at 8 and 12 (11.47% and 18.18%, respectively),
averaging 6.19%. Haplotrema accounted for 4.75% of the
total fauna.
ZONITIDAE
The 8 genera in our collections accounted for 38.52%
of the total fauna. Two genera, Mesomphix (13.44%)
and Ventridens (12.74%), are responsible for most of
the population, with Retinella (6.04%), Gastrodonta
(3.52%), and Striatura (1.40%) ranking third, fourth,
Vol. 12; No. 3
and fifth. The remaining 3 genera produced the following
relationships: Paravitrea (0.99%), Zonitoides (9.35%),
and Euconulus (0.17%). Similar data are presented for
individual species in Table 3.
Table 3
Percentage-Composition by Zonitid Species
in a Northern Kentucky Faunule
% of total
Species fauna
Euconulus chersinus (Say, 1821) 0.17
Retinella wheatleyi (BLAND, 1883) 1.64
Retinella indentata (Say, 1823) 0.05
Retinella carolinensis (CocKERELL, 1890) 0.88
Retinella cryptomphala (Cxapp, 1915) 3.46
Mesomphix inornatus (Say, 1821) 3.99
Mesomphix perlaevis (Pirssry, 1900) 6.92
Mesomphix vulgatus (H. B. Baker, 1933) 0.29
Mesomphix cupreus (RAFINESQUE, 1831) 1.93
Mesomphix capnodes (BINNEY, 1857) 0.29
Paravitrea species 0.05
Paravitrea multidentata (BINNEY, 1840) 0.11
Paravitrea placentula Prtspry, 1946 0.05
Paravitrea capsella (Goutp, 1851) 0.64
Paravitrea species a 0.05
Paravitrea species b 0.05
Gastrodonta interna (Say, 1822) S52.
Ventridens demissus (BINNEY, 1843) 12.74
Zonitoides nitidus (MULLER, 1774) 0.17
Zonitoides arboreus (Say, 1816) 0.17
Striatura ferrea Morse, 1864 1.40
Euconulus chersinus (Say, 1821)
Collecting sites: 7(2), 12(1)
Euconulus is not, according to our studies, a snail of
the dense forests. Our specimens were secured from
clumps of grass in clearings. ARCHER (1939) described
the species as being characteristic of the grassy slopes
of old gullies in Michigan.
Retinella wheatleyi (BLAND, 1883)
Collecting sites: 4(4), 5(6), 6(2), 7(10), 9(4), 10(1), 11(1)
Although widespread in the upland and middle stations,
Retinella wheatley: was not found to be the dominant
member of its genus. The snail was relatively more abun-
dant from station 4 through 7 (1.75% to 3.88% of the
total fauna) than from station 9, 10, and 11 (1.04% to
1.56%). The species lives primarily on the lower slopes
just above the stream, and on the flood plain, beneath
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Page 341
leaves, logs, rocks, and other moisture-conserving struc-
tures.
Retinella indentata (Say, 1823)
Collecting site: 1(1)
A single dead shell was collected at this site, and it
probably washed into the canyon from above.
Retinella carolinensis (CocKERELL, 1890)
Collecting sites: 7(1), 8(5), 9(6), 12(3)
Pirspry (1946) related that the type locality for this
species was the lower slopes of Roan Mountain (Great
Smokies), and he further stated that the species was prob-
ably found only on the lower slopes of protected valleys.
In this we concur. We did not secure specimens until
station 7, where the population was meagre (0.36% of
the total). The percentages at stations 8, 9, and 12 were
8.19, 4.44, and 5.46, respectively. An additional interesting
observation is, when Retinella carolinensis occurs sym-
patrically with R. cryptomphala, the former species tends
to replace the latter. At stations 8, 9, and 12, the popula-
tion percentages for R. cryptomphala were 3.27, 3.70,
and 3.63, respectively.
Retinella cryptomphala (Capp, 1915)
Collecting sites: 1(2), 2(4), 3(5), 4(1), 5(2), 6(6), 7(14), 8(2),
9(9), 10(1), 11(4), 12(2), 13(7)
Not only did this species occupy varying habitats — leaf
litter, decaying logs, clumps of grass, the undersides of
rocks — but it was found at all elevations, and from a
percentage comparison there seemed to be no differential
regarding preferences, other than the relationship men-
tioned above. The densest population, in relation to the
total fauna, occurred at station 3. However, this reflects
the decreased numbers of other species present rather than
an absolute increase in Retinella cryptomphala. Such local
variations are common in land snails, and are probably
responses to edaphic conditions.
Mesomphix inornatus (Say, 1821)
Collecting sites: 8(3), 9(14), 10(6), 11(14), 12(3), 13(8), 14(8)
Pi_spry’s statement concerning the habitat of this spe-
cies, 1. €., “moist, shaded slopes and ravines among
leaves and dead wood,” is supported by our findings.
Moreover, the absence of the species from the uplands
and head-valleys is striking. After its first appearance at
station 8, the species was present in dominant to co-dom-
inant numbers (4.91% to 14.58%), often sharing its po-
sition with the next species. We did not secure specimens
from the valley rim or upper slopes; all specimens were
found at the lower levels.
Page 342
Mesomphix perlaevis (Pirssry, 1900)
Collecting sites: 1(4), 2(6), 3(3), 4(3), 5(5), 6(13), 7(27), 8(10),
9(9), 10(1), 11(3), 12(5), 13(10), 14(12)
Found under dead leaves and decaying wood in the
middle to lower slopes at every station, and ranging from
1.56% to 16.39% of the fauna, this species has the widest
ecological distribution of any Mesomphix in the area. As
indicated above, in some areas it occupies co-dominant
status with M. inornatus. There did not appear to be pref-
erence for the small, heavily shaded valleys over the wider,
less heavily shaded ones, although the average percentage
of the total fauna from station 1 through 7 was 7.06
contrasted with 7.75 for stations 8 through 14.
Mesomphix vulgatus (H. B. Baker, 1933)
Collecting site: 9(5)
Because of the close-set spiral rows of microscopic pa-
pillae on the upper and lower surface of the last whorl,
these 5 specimens were tentatively identified as this spe-
cies. However, as HusricHt (1962b) pointed out, -diag-
noses based solely on shell characteristics in this group
are always open to question. Nevertheless, the shells are
nearly identical with ones from which the soft anatomy
was dissected. Mesomphix vulgatus is, with the next two
species, a member of third-order valley faunas, seldom
being found in the smaller, drier first and second order
systems. Wherever we have found the species accompanied
by M. perlaevis or M. inornatus, or both, it has always
been slightly to greatly outnumbered by them.
Mesomphix cupreus (RAFINESQUE, 1831)
Collecting sites: 2(5), 9(2), 10(5), 11(2), 12(2), 13(9), 14(8)
Regardless of the fair-sized sub-colony at station 2
(living in a pocket of deep humus against a fallen tulip
tree), Mesomphix cupreus characteristically is a middle
to lowland species, as evidenced by our percentage array
from stations 9 through 14: 1.48, 7.81, 2.08, 3.63, 8.03,
6.65, respectively. Pitspry’s (1946) habitat observations
coincide essentially with ours: “usually found partially
buried in damp humus, under a layer of dead leaves.”
Mesomphix capnodes (BinNEyY, 1857)
Collecting sites: 13(2), 14(3)
Our 5 specimens were removed from deep piles of
dead hardwood leaves on the west slope of the valley;
both sites were well shaded. Although we did not carry
our observations on into the lower Red River System, we
have collected the species from several loci since, always
in subordinate numbers. Mesomphix capnodes appears
to be another species of valleys of at least third-order
- magnitude.
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Vol. 12; No. 3
All of the Paravitrea specimens listed below were se-
cured from similar situations, i. e., from the interstices of
moist, deeply placed talus near the point where the slopes
leveled out into the floor of the valley. With the exception
of P. capsella (8.14%), none of the species was secured
in large numbers. The following percentages were ob-
tained: P multidentata (0.60% and 1.04%), P. placentula
(0.60%), P species a (0.60%) and P species b (1.75%).
Although too few specimens were found to allow deriva-
tion of ecological generalizations, the records are impor-
tant from the standpoint of succession (discussed below).
Paravitrea species
Collecting site: 4(1)
A single specimen, too immature to identify.
Paravitrea multidentata (BINNEY, 1840)
Collecting sites: 9(1), 11(1)
Paravitrea placentula lithodora Pitspry, 1946
Collecting site: 9(1)
Heretofore known only from Pine Mountain in Harlan
County, this form probably merits specific recognition.
Paravitrea capsella (Goutp, 1851)
Collecting site: 9(1)
The exact relationship of this species to its western
counterpart in the Ozarkian-Ouachitan region, Paravitrea
significans (BLAND, 1866), is quite hazy and in need of
detailed study.
Paravitrea new species? a
Collecting site: 2(1)
Shell small, pale yellowish, sub-shining; spire nearly
flat, with very shallow sutures minutely rebordered by
transparent shell material; 54 whorls, rather flat-sided,
curving gently to the sutures, the last slightly but defi-
nitely expanded; umbilicus deep, slightly lunate, nar-
row, the central portion about 9 times in diameter, but
expanding in the last whorl to about 5 times in diameter;
aperture deeper than wide, lunate; lip thin, simple; first
whorl smooth, on remaining ones growth striae raised,
irregular, becoming nearly regular on last two whorls;
spiral, incised sculpture minute above, more or less ob-
vious on otherwise smooth base; about one-third the way
in from aperture there is a transverse band of 5 very low,
contiguous tubercles.
Diameter 2.7 mm; height 1.3 mm; umbilical diameter
0.5 mm.
Deposited in the Field Museum of Natural History,
Chicago; No. FMNH 155477.
Vol. 12; No. 3
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Page 343
This form is most closely related to Paravitrea metallica
Husricut, 1963 (Husricut, 1963 c), being similar in
size, whorl configuration, and in possessing an expanded
last whorl. It differs from it in possessing a larger umbili-
cus, more regular sculpture, in having its aperture higher
than wide, and in having more and differently arranged
shell teeth. The teeth are arranged in one or two pairs in
P metallica, and the aperture is wider than high in that
species.
Paravitrea new species? b
Collecting site: 9(1)
Shell small, very pale tan, glistening; spire barely con-
vex, with shallow sutures, not re-bordered; whorls 534,
slowly expanding; upper periphery of body whorl slightly
angular, base well-rounded; umbilicus deep, well-like,
displaying all whorls, the central hole about 84 times
in diameter, expanding to about 4 times in the last
whorl; aperture only slightly wider than high, simple,
toothless ; nucleus of 14 whorls, smooth; sculpture of nearly
regular growth striae and radial grooves, crowded.
Diameter 2.5 mm; height 0.9mm; umbilical diameter
0.4 mm,
This species appears to be most nearly related to Para-
vitrea tantilla Husricut, 1963 (Husricut, 1963c) by
way of the shallow sutures, slowly increasing whorls, and
apertural and lip characters, but differs from that species
in possessing more regular sculpture, a larger umbilicus,
much smaller size (this form has nearly the same number
of whorls as P tantilla but is 1.2mm smaller), and by
lacking shell teeth.
Gastrodonta interna (Say, 1822)
Collecting sites: 1(11), 2(1), 4(2), 5(13), 6(2), 7(4), 9(10),
10(1), 12(1), 13(5)
Practically nothing has been published concerning the
ecology of this, one of the most distinctive genera of North
American snails. It lives on shaded slopes beneath dead
leaves, although it seems to avoid Rhododendron and
mountain laurel. At station 1 we found it abundantly
beneath white pine needles. ARCHER (1942) also found
the species associated with pine woods. There may be a
slight preference for smaller valleys over the larger ones.
The average composition-percentage in the upper valleys
(stations 1 through 7) was 4.35% (1.45% to 10.67%),
whereas in the lower ones (stations 9, 10, 12, 13) it was
3.46% (1.56% to 6.02%). However, sampling error
could account for this small difference.
Ventridens demissus (BINNEY, 1843)
Collecting sites: 1(28), 3(4), 4(30), 5(56), 6(33), 7(31), 8(24),
9(5), 10(4), 12(1)
Ventridens was absolutely the single most-abundant
species present in the study area from station 8 upward,
and is mainly responsible for placing the Zonitidae as the
second most abundant family. By station, the relative
percentages were (station numbers in parentheses) :
(DR 27e18i (3). 10:2551(4:)) 2951255 (5))) 35:89), (6) 28:94;
(7) 11.31, (8) 39.34, (9) 3.01, (10) 6.25, and (12) 1.81.
A sharp decrease in abundance was noted from station 9
downward, a trend we have also noted elsewhere. Our
specimens were collected mostly from the upper and
middle slopes, beneath dead leaves and logs. Such large
populations are common in the genus. Mayer (1965),
for example, reported one specimen of V. suppressus (Say,
1829) per square inch of log surface in Pennsylvania.
Zomitoides nitidus (Mu.ErR, 1774)
Collecting sites: 1(1), 2(2)
These specimens, representing 0.97% and 3.50% of the
total populations at the 2 stations respectively, were found
beneath dead leaves just above the stream margin. Pirs-
BRY (1946) indicated that the species was generally
found near water in lowland situations, never in upland
woods, and ARCHER’s (1939) observations were essentially
the same in Michigan.
Zonitoides arboreus (Say, 1816)
Collecting sites: 4(1), 7(2)
The specimen from station 4 was found under a dead
log on the flood plain, and those from station 7 under slabs
of sandstone about half-way up the slope. Although Zoni-
toides enjoys a much wider ecological range, up to at
least 10000 feet in the west, and down to sea level in
the south, it is never found far from moist situations.
Usually, the species is found in or around decaying wood.
At these 2 stations, the small numbers represented only
0.97% and 1.14% of the population, but in other locali-
ties we have it to constitute as much as 20%. The species
has a very erratic distribution within a given area.
Striatura ferrea Morse, 1864
Collecting sites: 1(8), 3(4), 4(2), 5(2), 8(2), 11(3), 13(3)
Following the same sequence as above, the composition
percentages at each station were: 7.76, 10.25, 1.94, 1.28,
3.27, 6.25, and 2.67. There does not appear to be definite
correlation between valley magnitude and the presence of
this small mollusk. Since the specimens all came from
the valley floors, under wet to damp leaves and decaying
wood, conditions near to or at dew point perhaps limit
the species’ distribution.
Page 344
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Vol. 12; No. 3
LIMACIDAE
Only a single species in this family was collected, repre-
senting only 0.70% of the total collection.
Deroceras reticulatum (MU.uer, 1774)
Collecting sites: 1(1), 4(10), 8(1)
All specimens of this widespread exotic species (GETZ,
1959) were found near the stream beneath decaying wood ;
the 10 at station 4 were collected from an old U.S. For-
estry marker. The percentages at the 3 stations were:
0.97, 9.70, and 1.63.
ENDODONTIDAE
Sixty-two specimens, or 3.61% of the total, were col-
lected. The genus Discus was most abundant, 2.75%,
with Anguispira (0.58%), Helicodiscus (0.17%), and
Punctum (0.11%) in less quantities. Since each genus,
except Discus, was represented by single species, the per-
centages were the same at the specific level. In Discus, D.
patulus represented 2.70% of the population, and D. bry-
anti only 0.05%.
Anguispira alternata (Say, 1816)
Collecting sites: 11(6), 12(2), 13(1), 14(1)
Our specimens are very definitely the lowland form
termed angulata by Pitspry (1948). They were collected
on the south slopes just above the stream, from beneath
decaying logs. This propensity for wood was also noted
by MacMiiian (1940). Percentage-wise, the species was
most abundant at station 11 (6.25%), and decreased in
abundance below that point, 3.63%, 0.89%, and 0.83%,
respectively. This possibly reflects the more open nature of
the last 3 stations.
Discus patulus (DeEsHAyYEs, 1830)
Collecting sites: 1(41), 2(1), 4(1), 5(3), 8(1), 9(11)
Another species with distinct decaying-woods affinity
(Husricut, 1963; Pirspry, 1948). There was a distinct
preference for the heavily shaded portions of the upper
valleys, especially the lower, moist slopes thereof. The
percentages by stations were: 22.33, 1.25, 0.97, 1.92,
1.63, and 8.14, respectively. The presence of the relatively
large population at station 9 probably was in response to
the dense shade and abundance of moist, decaying wood.
Discus bryanti (Harper, 1881)
Collecting site: 9(1)
Although this is characteristically a woodland species,
our siftings of forest floor debris did not disclose its pres-
ence at any of the upper stations. At the 2 lower stations,
where the population percentages ran 1.63 and 1.20, the
specimens were associated with decaying wood. From past
observations, the senior author feels that this and other
Helicodiscus and related species tend to avoid Rhodo-
dendron. Consequently, since H. parallelus has been found
at 5000 feet elevation in Tennessee (Pmssry, 1948), the
apparent restriction to the lower valleys may be correlated
with the marked decrease in Rhododendron at those sta-
tions. Bascu et al. (1961), however, found the species
more common on flood plains.
Punctum minutissimum (Lr, 1841)
Collecting site: 7(2)
Our specimens were found beneath a deep pile of de-
caying hardwood leaves. According to Morse (in Pits-
BRY, 1948), this minute species prefers the “rotten bark
of beech trees,” a common tree in the study area, although
somewhat scattered. Our scanty records, of course, did
not allow us to corroborate Morse’s statement.
PHILOMYCIDAE
The two genera and 4 species of these rather typical
eastern slugs made up 2.16% of the total collection, with
Pallifera accounting for 1.40%, and Philomycus for
0.76%. By species, the following data were obtained:
Philomycus carolinianus 0.17%, P. flexuolaris 0.58%,
Pallifera dorsalis 1.18%, and Pallifera fosteri 0.23%.
Philomycus carolinianus (Bosc, 1802)
Collecting sites: 1(1), 10(2)
Only collected from the space between the bark and
trunk of decaying hardwood logs on the flood plains and
lower slopes, our records corroborate BRANSON’s (1962)
conclusion that this is a flood plains species. It probably
occurs throughout the system. At the 2 localities, Philo-
mycus carolinianus accounted for 0.97% and 3.12% of
the fauna. INcram (1949) found this slug in New York
beech-hemlock associations, but observed that it avoided
hemlock logs.
Philomycus flexuolaris (RAFINESQUE, 1820)
Collecting sites: 4(4), 8(2), 9(3), 12(1)
Long considered a subspecies of Philomycus carolini-
anus (Pitspry, 1948, and others), Husricut (1951)
demonstrated the distinctness of this large brownish slug.
Ecologically it is also distinct, since the preferred habitat
is beneath stones and in rock fissures in the upland slopes.
All of our specimens were found beneath logs, lying on
sand, stone, or under rocks. In the station order shown
above, the percentages ran 3.88, 3.27, 2.22, and 1.81.
Vol. 12; No. 3
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Page 345
Pallifera dorsalis (BINNEY, 1842)
Collecting sites: 3(1), 6(3), 7(2), 10(1), 12(6), 13(4), 14(3)
Although widespread in the valley system, this form
had a rather decided preference for the lower stations. It
was found on the lower slopes and on the flood plains
beneath decaying leaves, along the edges of logs, and
beneath stones. DimELow (1962) found Pallifera dorsalis
common in Nova Scotian deciduous leaf litter. The per-
centages were: 2.56, 2.63, 1,14, 1.56, 10.90, 3.57, and
2.50.
Pallifera fosteri F.C. Baker, 1939
Collecting sites: 3(1), 7(2), 10(1)
The external and internal anatomy of these specimens
was in essential agreement with the findings of Grimm
(1961) and Wess (1952). From past observations (BRAN-
SON, 1962), it is apparent that this form is strictly a
species of very moist flood plains. All of our specimens
were removed from decaying logs in mud flats.
PUPILLIDAE
Only 2 specimens of one species were collected. Gastro-
copta is not in general a well-adapted mountain genus,
although certain sections of the genus are. The species dis-
cussed below, for example, seems to be absent from the
higher parts of the Allegheny Mountains (Pitspry, 1948),
and Cuan & Kemp (1930) reported the species as only
locally abundant in an Indiana locality. At lower eleva-
tions, G. contracta prefers mesic situations under dead
woods and leaf mold (Bascu e¢ al., 1961). In our
study, Gastrocopta comprised only 0.11% of the total
fauna.
Gastrocopta contracta (Say, 1822)
Collecting sites: 2(1), 9(1)
The percentage composition at station 2 was 1.75, at
station 9 it was 0.74.
CIONELLIDAE
This family is represented in North America by a single
widespread species, Cionella lubrica (MULLER). In this
study it represented 1.05% of the fauna, all of our speci-
mens being secured from the moist, heavily-shaded spots
at each station, beneath thick layers of damp to wet
decaying leaves.
Cionella lubrica morseana DoHERTY, 1878
Collecting sites: 1(1), 2(4), 7(2), 8(1), 9(5), 10(1), 12(1)
_ The following percentages were calculated for the above
Stations O:9/ee/Oly Ll4 63.0 370 leo On lol.
AQUATIC SPECIES
It has often been observed that slightly alkaline waters
are more productive of mollusks than soft ones. However,
very few reports present chemical characteristics of the
water from which gastropods were collected. SHoup
(1943), studying various streams in the Obey River
Drainage, Tennessee, found that streams flowing over
sandstones of the Lee Formation were in general poor
in bicarbonates, some of them as low as 2 to 6 parts per
million. According to his findings, streams with bicarbon-
ates below 20 ppm possessed depauperate mollusk faunas.
In the discussion which follows, it will be noted that
mollusks were entirely lacking at all of the upland stream
stations, not appearing before station 11. A glance at
Table 1 will reveal very low bicarbonate in the crystal
clear water until station 11, where the stream leaves the
Lee Formation, and where the bicarbonate reaches a
level above 40 ppm.
There are, of course, other considerations. Many aquatic
snails are invariably associated with rooted vegetation,
such as Campeloma (A.uson, 1942; Bovpyerc, 1952),
Lioplax, and certain species of Physa (BickeL, 1965).
Since the water in our study area is subject to much
fluctuation in depth, and probably also because of low
carbonate content, rooted plants are very scarce to ab-
sent. In other words, the type of marginal habitat re-
quired by amnicolid and physid snails is lacking in the
headwater situations, but becomes progressively more
available downstream.
In our study 4 families of aquatic snails were encoun-
tered, although the first form discussed is actually amphi-
bious.
LYMNAEIDAE
Lymnaea humilis Say, 1822 (= L. dalli F C. Baker),
the single species (7 specimens) collected by us at station
11, comprised 0.41% of the total fauna, and 7.29% of
the fauna at the collecting site. As indicated above, this is
an amphibious form; our specimens were found crawling
on vertical sandstone some 5 feet above the water. This
is apparently common, as Baker (1911) indicated that
the species was “generally out of water on sticks and
stones or mud flats.”
PHYSIDAE
Only a single species in this nearly ubiquitous family was
secured, representing 0.76% of the total collection.
Page 346
THE VELIGER
Vol. 12; No. 3
Physa integra HALDEMAN, 1843
Collecting sites: 13(5), 14(8)
The local percentages were 4.46 and 6.67, respectively.
As already mentioned, BickeL (1965) has demonstrated
this species’ dependency upon rooted vegetation. Since
the upper stations had little vegetation, its absence there
could be expected. Our specimens were secured in back-
water situations on decaying leaves and Potamogeton.
Outside the study area, downstream in Red River, the
species becomes abundant on Dianthera, where it is
associated also with Campeloma.
ANCYLIDAE
The ecology of this very interesting family, which ac-
counted for 0.52% of the mollusks found by us, remains
sketchy. Two species, in separate genera, were secured.
Laevapex fuscus (C. B. Apams, 1840)
Collecting site: 14(1)
Characteristically a species of lakes and slow-flowing
rivers, our single specimen (0.83% of the local fauna) was
removed from a dead leaf in a side-pool.
Rhodachmea elatior (ANTHONY, 1855)
Collecting site: 14(8)
These specimens, comprising 6.67% of the local fauna
and 0.40% of the total collection, are the first collected in
Kentucky since ANTHONY described the species from
Green River. This is a rather important discovery, since
Bascu (1963) indicated that the species was known def-
initely only from the Cahaba River in Alabama. Our spe-
cimens were found on stones in a swift riffle.
AMNICOLIDAE
Somatogyrus subglobosus (Say, 1825)
The single specimen of Somatogyrus subglobosus (Say,
1825) found at station 14 constituted 0.05% of the total
collection and 0.83% of the local fauna. It was associated
with the Rhodachmea.
DISCUSSION anp CONCLUSIONS
Longitudinal succession in stream organisms, from head-
waters downward, is a well-known principle (Opum,
1959), one which KuEHNE (1962) has correlated with
Horton’s (1945) drainage classification as modified by
STRAHLER (1954, 1957). Horton’s system is based on
stream branching, and intermittent or permanent head-
water streams are classified as Order 1. The union of 2
equal-ranked streams forms one of the next highest order,
but the rank of the stream is not increased by the entrance
of one of lower order than itself, KuEHNE’s (op. cit.) work
showed a progressive increase in the average number of
fish species present with an increase in stream order, and
he suggested this relationship reflected adaptations to
local conditions.
With these studies in mind, we decided to sample mol-
lusk populations to test the hypothesis that valleys them-
selves could be ranked by order, which in turn would be
reflected in the terrestrial fauna. The correlation with
stream order is obvious in the aquatic forms (see above).
In terrestrial species there also seems to be a definite
correlation between valley order and number of species
present (Figure 3). The average number of species at the
Station
123 4 5 6 7 8 9 10 11 12 13 14
Number of Species
oO
Figure 3
Graph showing Species Abundance per Station.
Average for upper 7 stations: 17.3; average for lower 7 stations: 20.6.
upper 7 stations was 17.3, contrasted with 20.6 at the
lower 7. Figure 4 shows the distribution of all species
collected, thereby demonstrating longitudinal succession.
None of the species was restricted to order 1 or order 2
valleys, but 4 species, Mesodon thyroidus, Mesomphix
inornatus, Mesomphix cupreus, and Retinella carolin-
ensis, were found only in order 3 and 4 valleys. Several of
the other species, such as some Paravitrea, may also be
restricted to the larger systems.
Another aspect of longitudinal succession observed was
that of relative abundance. Roughly, 4 main patterns were
observed (Figure 5): most abundant in order 1 and 2
valleys with decreasing abundance downgrade (Ventri-
Vol. 12; No. 3 THE VELIGER Page 347
n
OO OO|~s
°
i=]
a
—
Oo
—
—
Stenotrema evardsi
il 2
© ©
Stenotrema stenotrema ©) ©
OQ ©
OO] &
OO
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Stenotrema angellum
Stenotrema barbatum
Stenotrema leat
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Mesodon thyroidus
Mesodon zaletus
Mesodon appressus
Mesodon sayanus
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Mesodon ruglet
O|OOO
O|OO0O
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Triodopsis tridentata
Triodopsis fraudulenta
Triodopsis denotata
Triodopsis albolabris
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©m|COCOO||©
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OO
O
OO
OO
O|\OO O© |OO©
O
Euconulus
Retinella wheatleyi CO. O ©
Retinella indentata
Retinella carolinensis
Retinella cryptomphala
© |©OOO C|OO©
@)
e)
O
O
O
©
Mesomphix inornatus
Mesomphix perlaevis CoO O OO © ©O
Mesomphix vulgatus
Mesomphix cupreus
Mesomphix capnodes
© OCG COOLO OClOOO©
OO/|OO
QO OO|©
OQ CO \W©
© OO|O©
OO OO|©
COO O©
O
Paravitrea placentula
Paravitrea multidentata
Paravitrea capsella
Paravitrea lafuzi
Paravitrea sceada
O
Gastrodonta
Ventridens
Zonitoides nitidus
Zonitoides arboreus
Striatura
OO O}] ©
O
OO
OO
OO
OO
O
OO
OO
O|O
Derocerus
Anguispira
Discus patulus
Discus bryanti
Punctum O
Philomycus carolinianus
Philomycus flexuolaris O @e@®
Pallifera dorsalis
Pallifera fosteri
Gastrocopta
Cionella
Lymnaea O
Physa @
Laevapex
Rhodachmea
Somatogyrus
© ClOOOOl
O
O
O
CO OO
OO
O
O;O
OO
O
O
OC! COOO
O
O
O
OOOO
Figure 4
Graph showing Occurrence.of Mollusks at Collecting Stations.
Station
3G fy
I 2
Ventridens
25
Number of Specimens
Mesodon
evardst
Mesodon
Sayanus
Figure 5
Graph showing Distribution by Numbers in Four Molluscan Species.
dens), most abundant in order 2 and order 3 valleys (Ste-
notrema evardsi), most abundant in order 4 valleys (Me-
sodon sayanus), and more or less abundant throughout
the system (Mesodon appressus). Other species could
have been selected, but these are representative. Some of
the sporadic variation may be the result of local differ-
ences in edaphic conditions (BurcH, 1955) and, indirect-
ly, the occurrence of favorable plant associations (BURCH,
19577)
A final, and perhaps more important, observation is
illustrated by Figures 6 and 7. It appears that when 2
snail species with similar habitat requirements occur to-
gether sympathically one or the other’s population is
held in abeyance, probably according to which species is
favored by the particular environment. In the case of
Mesodon appressus, which we have already indicated
_ produces its largest populations in the uplands, versus M.
sayanus (Figure 6), it can be seen that M. sayanus is
THE VELIGER
7 8 9 TO Re oe Ue} wi
Vol. 12; No. 3
Station
1 2 3 4 5 6 7 8 g 10 11 12 13 14
30
ew
°
Te
3
% 20 Mesodon
o — sayanus
8
3 |
Hi
§ Mesodon
5 10 — appressus
a
5
Figure 6
Graph showing Station Percentages for Mesodon appressus
and Mesodon sayanus.
held to relatively small populations in the uplands, where-
as there is a tendency to reversal in order 3 and order 4
valleys. In Mesomphix perlaevis versus M. inornatus the
contrast is even more striking (Figure 7). These compe-
titive relationships need intensive study.
Comparisons of disturbed versus undisturbed areas
showed no positive correlations.
8 9 10 11 12 13 14
20
~
ao
Mesomphix
inornatus
Mesomphix
perlaevis
o
Percen Station Collection
C)
Figure 7
Graph showing Station Percentages for Mesomphix perlaevis
and Mesomphix inornatus.
——————————————————— a i ea ee
Vol. 12; No. 3
THE VELIGER
Page 349
SUMMARY
1. Fourteen randomly selected stations, located in Wolfe
and Powell Counties, Northern Kentucky, were sys-
tematically searched for terrestrial and aquatic mol-
lusks, the habitat conditions at each station were
detailed, and information on plant associations and
water chemistry secured.
2. Twelve families, 26 genera, and 52 species, two ap-
parently new, were collected.
3. The family Polygyridae is the dominant taxon present
in the study area, 47.26% of the total fauna. It is
followed by the Zonitidae, 38.52%. The remaining
14.22% of the fauna is unequally divided between the
other 10 families.
4. The percentage-composition of the total fauna was
calculated for each genus, showing Mesodon (23.67%),
Stenotrema (15.03%), Mesomphix (13.44%), and
Ventridens (12.74%) to be the dominant ones.
5. The number of each species collected per station is
presented; percentages were calculated as regards the
total fauna and the faunas at each station. Mesodon
ruglei (10.74%) and Ventridens demissus (12.74%)
are the two dominant species in this region.
6. Observations on habitat and ecology are presented in
discussions of individual species.
7. Analysis of ecological distribution demonstrates a
positive correlation between the number of species
present and valley magnitude. No species were restric-
ted to valleys of first or second order, but several were
found only in order 3 or order 4 valleys.
8. Correlated with low concentrations of calcium car-
bonate in order 1, 2, and 3 streams, aquatic mollusks
are restricted to order 4 streams in the study area.
9. Relative abundance observations disclosed 4 main
patterns: (a) greatest abundance in order 1 and 2
valleys, decreasing in abundance downgrade; (b) great-
est abundance in order 2 and 3 valleys, decreasing up-
and downgrade; (c) greatest abundance in order 4
valleys, decrease upgrade; and (d) abundant through-
out the system. Examples of each type of distribution
are included.
10. Sympatric species with similar ecological require-
ments, according to local environmental conditions,
mutually affect populations, either by suppressing an
associated species or being suppressed by it.
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1911. |The Lymnaeidae of North and Middle America.
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1963. A review of the Recent freshwater limpet snails of
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1965. The role of aquatic plants and submerged structures in
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952% Ecological aspects of the dispersal of the snail Campe-
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BRANSON, BRANLEY ALLAN
1962. The slugs (Gastropoda : Pulmonata) of Oklahoma and
Kansas with new records. Trans. Kansas Acad. Sci. 65:
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BRANSON, BRANLEY ALLAN & DoNALD LEE BaTcH
1968. Ecological study on valley-forest spiders from Northern
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1969. Valley centipedes (Chilopoda : Symphyla) from North-
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Burcu, JoHN BayArp
1955. Some ecological factors of the soil affecting the distribu-
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Caun, Arvin Ropert & Jack T. Kemp
1930. The terrestrial mollusks of Turkey Run State Park, In-
diana. Trans. Illinois State Acad. Sci. 22: 250 - 262
CLENCH, WILLIAM JAMES
1962a. A catalogue of the Viviparidae of North America with
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lusca, Mus. Comp. Zool. Harvard 2: 288
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THE VELIGER
Vol. 12; No. 3
1926. Some notes and a list of shells of Rio, Kentucky.
The Nautilus 40 (1): 7-12; (2): 65-67
CLENcH, WILLIAM JAMES & RuTH Drxon TuRNER
1955. The North American genus Lioplax in the family Vivi-
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Conkuin, James ELWIN
1957. ‘The larger land snails of Sleepy Hollow, Kentucky.
The Nautilus 71 (1): 10-11
DimeELow, E. J.
1962. On the biology of some mollusks from a Nova Scotian
deciduous wood. The Nautilus 76: 49 - 51
Getz, LowE tu L.
1959. Notes on the ecology of slugs: Arion circumscriptus,
Deroceras reticulatum and D. laeve. Am. Midl. Nat.
61: 485 - 498
GoopricH, CALVIN
1921. Three new species of Pleuroceridae. Occ. Pap. Mus.
Zool. Univ. Michigan 91: 1-5
1929. The pleurocerid fauna of the Ohio. The Nautilus
43(1):1-17
1934a. Studies of the gastropod family Pleuroceridae. I
Occ. Pap. Mus. Zool. Univ. Michigan 286: 1 - 17
1934b. Studies of the gastropod family Pleuroceridae. ITI
Occ. Pap. Mus. Zool. Univ. Michigan 295: 1-6
1934c. Studies of the gastropod family Pleuroceridae. III
Occ. PapersMus. Zool. Univ. Michigan 300: 1 - 11
1937. Studies of the gastropod family Pleuroceridae. VI
Occ. Pap. Mus. Zool. Univ. Michigan 347; 1 - 12
1938. Studies of the gastropod family Pleuroceridae. VII
Occ. Pap. Mus. Zool. Univ. Michigan 376: 1 - 12
1940. The Pleuroceridae of the Ohio River drainage system.
Occ. Pap. Mus. Zool. Univ. Michigan 417: 1 - 21
1941. Studies of the gastropod family Pleuroceridae. VIII
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Grimm, FE Wayne
1961. Pallifera fosteri, with P. megaphallica, new. The
Nautilus 74 (3): 102-105; 2 text figs.
Horton, Rosert E,
1945. _Erosional development of streams and their drainage
basins; hydrophysical approach to quantitative morphology.
Bull. Geogr. Soc. Amer. 56: 275 - 370
Hupricut, LESLIE
1950. Mesodon andrewsae normalis (Pits.) in Kentucky.
The Nautilus 63 (3): 106
1951. The Limacidae and Philomycidae of Pittsylvania Coun-
ty, Virginia. The Nautilus 65 (1): 20-21
1958a. New species of land snails from the Eastern United
States. Trans. Kentucky Acad. Sci. 19: 70 - 76
1958b. Quickella vermeta and Succinea indiana. The
Nautilus 72 (2): 60-61
1958c. New species of land snails from the Eastern United
States. Trans. Kentucky Acad. Sci. 19: 70 - 76
1960. The cave snail, Carychium stygium CAL.
Kentucky Acad. Sci. 21: 35 - 38
Trans.
1962a. New species of Helicodiscus from the Eastern United
States. The Nautilus 75 (3): 102-107; plts. 7-9; 2 text
figs. -
1962b. Mesomphix vulgatus and its allies. The Nautilus
76 (1): 1-3
1963a. Carychium exile and Carychium exiguum. The
Nautilus 76: 108
1963b. New species of Hydrobiidae. Nautilus 76: 138 - 140
1963c. Four new species of Paravitrea. The Nautilus 76:
140 - 142
1964a. The bidentate species of Ventridens (Stylommatophora:
Zonitidae) . Malacologia 1: 417 - 426
1964b. Land snails from the caves of Kentucky, Tennessee, and
Alabama. Bull. Nat. Speleol. Soc. 26: 33 - 34
INGRAM, WILLIAM Marcus
1949. Natural history observations on Philomycus carolinianus
(Bosc). The Nautilus 62 (3): 86-90
Kaptan, MicHatEL F « W. L. MIncKLEY
1960. Land snails from the Doe Creek area, Meade County,
Kentucky. The Nautilus 74 (2): 62-65
KuEHNE, RoseErt A.
1962. A classification of streams, illustrated by fish distribu-
tion in an eastern Kentucky creek. Ecology 43: 608 - 614
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1949. Determination of color of turbid waters. Anal.
Chem. 21: 276
MacMiLian, Gorpon KuTcHKA
1940. A monographic study of the snails of the genera Angui-
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1965. Brief census of log-associated snails in Berks county,
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1959. Fundamentals of Ecology.
Philadelphia, 546 pp.
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1965. Standard methods for the examination of water and
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Public Health Assoc., Inc., New York, 769 pp.
Pitspry. Henry AUGUSTUS
1939. Land Mollusca of North America, North of Mexico.
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1959. + Mollusks of the Salt River, Kentucky. The Nautilus
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1943. Distribution of fresh-water gastropods in relation to
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1957. Quantitative geomorphology of erosional landscapes.
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podia 1: 6-7
Vol. 12; No. 3
THE VELIGER
Page 351
A New Species of Muricopsis from West Mexico
BY
GEORGE E. RADWIN
AND
ANTHONY D’ATTILIO
San Diego Natural History Museum, San Diego, California 92112
(Plate 52; 4 Text figures)
IN THE COURSE OF our continuing study of the Muricidae
of the Panamic province of the eastern Pacific we have
frequently encountered an apparently undescribed species
of Muricopsis from Jalisco state, West Mexico. Almost two
years ago the junior author was first shown specimens of
this species by Mrs. Ruth Purdy of San Diego, California.
Since then, many specimens have been generously made
available to us for examination. The number of specimens
in private and museum collections is surprisingly large and
is indicative that the species is at least moderately com-
mon at certain points in the region between Banderas Bay
(Puerto Vallarta) and Manzanillo.
ABBREVIATIONS USED
SDNHS - San Diego Natural History Society
LACM - Los Angeles County Museum of Natural
History
AMNH - American Museum of Natural History
Muricopsis Bucquoy & DAUTZENBERG,
1882
Muricidea Morcu, 1852, p. 95, non Swainson,
1840; type species: Murex magellanicus GMELIN,
1791 (=M. geversianus Patuas, 1774) by OD
Muricopsis Bucguoy & DAUTZENBERG, in Buc-
guoy, DAUTZENBERG & Do.tiFus, 1882, p. 19; type
species: Murex blainvillet PAYRAUDEAU, 1862, by OD
Remarks: The genus Muricopsis was originally erected
for mollusks with small to moderate-sized, spinose or tu-
berculate shells, znd with strong denticulation on the inner
surface of the apertural lip. Color and degree of sculp-
ture vary even within a single species as is noted by
Bucqguoy & DAUTZENBERG.
The following nominal species from the eastern Pacific
are apparently referable to Muricopsis.
1. Murex armatus A. Apams, 1854, p. 71, Gulf of Cali-
fornia, Mus. Cuming, (= Muricidea squamulata Car-
PENTER, 1865), holotype, BM(NH) no. 1954-4.15.10-12
(see Keen, 1958, fig. 367).
2. Muricopsis zeteki HERTLEIN & STRONG, 1951, p. 85,
Panama City, Panama, (== Murex aculeatus Woop, 1828,
non Lamarck, 1822) (= Murex dubius Sowersy, 1841,
non Dittwyn, i817), holotype (of M. aculeatus): ?>BM
(NH), see Dance, 1966, p. 305.
3. Murex pauxillus A. ApaMs, 1854, p. 71, Gulf of Cali-
fornia, Mus. Cuming, holotype: BM(NH).
4. Muricopsis jaliscoensis, new species, described below,
from Jalisco, West Mexico.
The genus Muricopsis has an apparently long and wide-
spread fossil distribution. According to WENz (1941),
fossil species are known from most world areas; the ear-
liest, reported from the Paleocene of Europe. These in-
clude: Europe (M. multistriatus DESHAvES), Indo-Paci-
fic, North America (M. spinulosa Hettrrin, M. aldrichi
CossMANN) and Australia (M. alveolatus TATE).
Muricops’s blainvillei (PAYRAUDEAU, 1826)
(Plate 52, Figure 3)
Murex blainvillii PayRAUDEAU, 1826, p. 149; plt. 7,
figs. 17, 18; L’fle de Corse (Corsica).
Murex cristatus Broccut; Risso, 1826, vol. 4, p. 191.
Cancellaria blainvillt PayRAUDEAU; BLAINVILLE,
1826, p. 139; plt. 5, fig. 4.
Page 352
THE VELIGER
Vol. 12; No. 3
Figure 1
Muricopsis blainville: (PAYRAUDEAU, 1826)
one lateral and one central tooth from the radula
(SDNHS no. 51308)
Murex pliciferus Brvona-BERNARDI, 1832, p. 22; plt.
3, fig. 10.
Murex (Muricidea) blainvillit, PayrAUDEAU; H. & A.
Apams, vol. 1, p. 75.
Murex (Ocinebra) blainvillii PayrAuDEAU; MonrTE-
ROSATO, 1878, p. 40.
Murex (Muricopsis) blainvillei PAyRAUDEAU; Buc-
Quoy & DAUTZENBERG, 1882, vol. 1, p. 19; Roussillon,
France.
Distribution: Mediterranean Sea, Atlantic coast of Por-
tugal, Spain and North Africa, as well as the Island of
Madeira. Also reported, under the name of Murex crista-
tus Broccut, 1814 from the Pliocene of Italy.
Description of the radula: Each transverse radular row
has a single rachidian tooth, flanked on each side by a
simple, sickle-shaped lateral tooth (Text figure 1). The
base of the rachidian tooth is roughly rectangular with
very prominent, downward pointing projections on each
end of the rectangle, and 5 sharp cusps. The lateral cusps
are quite long and sharp, the intermediate cusps are as
sharp, about % as long, and are positioned closer to the
lateral cusps than to the central cusp. The central cusp
is the longest and strongest of the 5, with a prominent,
blade-like highlight along its leading edge. It is borne on
a cowl-like structure and projects far in advance of the
remainder of the tooth. It is not as down-hooked as is
the case in Muzicopsis jaliscoensis and in some other
species of Muricopsis. As a result of this great forward
extension of the central cusp, it is virtually impossible,
under high magnification, to bring it into focus simulta-
neously with the remainder of the cusps.
Remarks: The type locality of Muricopsis blainvillei is
Corsica in the northwestern Mediterranean. For many
years, another name has persistently competed with M.
blainvillet for primary recognition. Murex cristatus was
first described by Broccut (1814) for a fossil species from
Explanation of Plate 52
Figure 1: Muricopsis zeteki HERTLEIN & STRONG, 1951, SDNHS no.
50821, Bahia Coastocomate, Jalisco, Mexico; length 21.2 mm;
width 13mm
Figure 2: Muricopsis armatus (A. Apams, 1854), SDNHS no.
22740, La Paz, Baja California del Sur, Mexico; length 32 mm;
width 18 mm :
- Figure 3: Muricopsis blainvillei (PayRAUDEAU, 1826), SDNHS no.
51308, Livorno, Mare Tirreno, Italy; length 24 mm; width 12mm
Figure 4: Muricopsis jaliscoensis RADWIN & D’ATTILIO, spec. nov.
holotype, SDNHS no. 51251, Bahia Coastocomate, Jalisco, Mexico;
length 26.5 mm; width 13.8 mm
Figure 5: Muricopsis jaliscoensis RADwIN & D’ATTILIO, spec. nov.
paratype, SDNHS no. 51250, Bahia Coastocomate, Jalisco, Mexico;
length 23.3 mm; width 13mm
Figure 6: Muricopsis jaliscoensis RADWIN & D’ArTILIO, spec. nov.
protoconch, greatly enlarged
THE VELIGER, Vol. 12, No. 3
[Rapwin & D’Artixio] Plate 52
Figure 1 Figure 2 Figure 3
Figure 4 Figure 5 Figure 6
Vol. 12; No. 3
the Pliocene of Italy. Although this name has often been
used for M. blainvillei, we have been unable to come to a
firm conclusion concerning its status. We have never seen
a bona fide Recent specimen of Murex cristatus and, there-
fore, any statement on this matter would enter the realm
of conjecture. If Murex cristatus were to be shown to be
the same as Muricopsis blainvillei, the former name
would, of course, have priority and the latter would be
considered a junior synonym.
We have not seen the type of Murex blainvillei, said by
Dance (1966, p. 297) to be in the Paris Museum, but
there is little doubt about the identity of this species,
figured many times in the literature (PAYRAUDEAU, 1826;
GrancER, 1884; Locarp, 1892).
Muricopsis jaliscoensis Rapwin & D’ATTILIO, spec. nov.
(Plate 52, Figures 4, 5)
Description: Shell light brown; length 26.5 mm; 6 whorls
excluding the nucleus; spire high, with 5 spinose varices
and sculptured with finely scabrous striae throughout. The
nucleus of an immature paratype has 2 whorls; the first
whorl is somewhat flat and slightly depressed; the second
nuclear whorl is distinctly tabulate (Plate 52, Figure 6).
Axial sculpture on the spire consists of conical tubercles at
the shoulder, extending axially to preceding whorls as
low ridges; areas between these low ridges equal in
width and concave; suture very shallow and undulate.
Spiral sculpture on the body whorl consists of primary,
secondary, and tertiary elements. There are 5 primary
cords, 4 equidistant cords and a fifth, more distant one
on the canal. Each of these ends in a short varical spine;
spines increase in length toward the final varix. The vari-
cal edge is obliterated except on the last 3 or 4 varices.
Between the primary cords are 1 or 2 secondary cords, also
ending in tiny spines. Interspersed between the primary
and secondary elements are numerous fine tertiary threads,
imbricated with close-set scales, present also on the vari-
cal spines. In addition, the shoulder has about 10 spiral
cords of secondary strength. The spiral cords on the spire
are the same size and development as those on the body
whorl. The color of the entire skell is an even tone of
yellow-brown. The aperture is of moderate size; situated
below the median area of the shell; is white to bluish
white; ovate; with a posterior anal groove; the inner sur-
face of the apertural lip has 7 denticles, arranged as
follows: a large knob-like one near the anal groove; be-
low this, occupying the remainder of the lip are 6 den-
ticles; the center 4 set more closely together, the remaining
2 somewhat more distant, 1 on each side of the central 4.
The columellar edge is slightly erect anteriorly. On the
THE VELIGER
Page 353
columella, directly above the siphonal canal are 3 conti-
guous denticles in a posterior-anterior orientation, the
posterior one most prominent. The siphonal canal is mod-
erately long, narrowly open and recurved; the siphonal
fasciole bears the short distal portions of 3 previous canals.
Description of the radula: Each transverse radular row
has a single rachidian tooth, flanked on each side by a
simple, sickle-like lateral tooth (Text figure 2). The base
of the rachidian tooth is roughly rectangular with a prom-
inent, downward pointing projection at each end of the
rectangle and 5 sharp cusps. The lateral cusps are moder-
ately long and sharp. The intermediate cusps are as sharp
but only half as long and are positioned closer to the
lateral cusp than to the central. The central cusp is un-
usually large and strong and is borne on a cowl-like
structure. This structure enables it to project further out-
ward and downward than any of the other cusps. Because
of the great distance that this cusp projects in advance
of the others, it is impossible, at the desired high magni-
fication, to bring it into focus simultaneously with the
remainder of the cusps.
Measurements: holotype — length, 26.5 mm; width, 13.8
mm; largest paratype — length 27.3 mm; width, 16.5
mm (lacking protoconch) ; smallest mature paratype —
length, 22.2 mm; width, 12.0 mm.
Type locality: Bahia Coastocomate, Jalisco, Mexico, in
20 - 70 feet, Lawrence E. Thomas, January, 1969 (holo-
type, see Plate 52, Figure 4; and 4 paratypes).
Type depositories: holotype, SDNHS no. 51251; 4 para-
types, SDNHS no. 51250, January, 1969 (Plate 52, Figure
5), both from type locality; 3 paratypes, SDNHS no.
51015, March 1, 1969 (specimen from which figured
radula was extracted), Puerto Vallarta, Jalisco, Mexico,
2 mature, 1 immature; 1 paratype, SDNHS no. 51285,
October 19, 1968, Coastocomate, Jalisco, Mexico, mature;
22 paratypes, 5 mature, 17 immature, LACM no. 65-15,
March 22, 1965, Los Arcos, Banderas Bay, Jalisco, Mexi-
co; 30 paratypes, 18 mature, 12 immature, LACM no.
65-14, March 21, 1965, Tres Marietas Islands, Banderas
Bay, Jalisco, Mexico; 5 paratypes, all mature, AMNH no.
153349, February 1, 1969, Barra de Navidad, Jalisco,
Mexico (dead specimens) .
Remarks: The holotype is a fresh, live-taken specimen,
as are all but 5 of the paratypes (see above). The 5 para-
types deposited at the American Musuem of Natural His-
tory are dead-collected but rather fresh.
From a zoogeographic viewpoint the distribution of this
species is narrowly restricted, on the basis of our present
knowledge. It is found mainly in the area between Ban-
deras Bay (Puerto Vallarta) and Manzanillo. This portion
Page 354
THE VELIGER
Vol. 12; No. 3
a ee es,
So ey
Figure 2
Muricopsis jaliscoensis Rapwin & D’ATTILIO, spec. nov.
one lateral and one central tooth from the radula of a paratype
(SDNHS no. 51015)
of Mexico has not, apparently, been sufficiently explored
for its molluscan species. Although the new species is not
rare, and found in fairly shallow water, the lack of collect-
ing in this relatively small area of the West Mexican coast
may account for its not being discovered until the last
few years. It is most closely allied in size and form to
Muricopsis zeteki HERTLEIN & STRONG, 1951 (Plate 52,
Figure 1), a widespread species occurring from the Gulf
of California to Ecuador and Galapagos. Muricopsis ze-
teki differs, however, in its more slender overall appear-
ance and its more spinose character. In the area in which
both species occur, the spines and primary spiral cords
of M. zeteki are whitish, the interspaces between the
spines are dark brown to black, and the remainder of the
shell is yellow-tan. One other species, M. armatus (A.
ApaMS, 1854) (Plate 52, Figure 2) needs to be considered
in this context. The distribution of M. armatus is from
the upper portion of the Gulf of California to Mazatlan,
Mexico, with dubious records from Nicaragua and Pana-
ma. Its distribution does not seem to overlap that of M.
jaliscoensis. Muricopsis armatus is whitish to pale yellow
- or brown in color; it is larger (18 - 47 mm) and in areas
of optimal conditions the spines are long and sharp. The
shoulder is well defined. The inner surface of the apertural
lip has 5 to 7 denticles, depending on the size of the indi-
vidual. On the anterior columellar edge there are 2
denticles; 1 at the mouth of the aperture and 1 recessed
more deeply.
In the aperture of all these species, one may find a
quick diagnostic feature for their separation. Muricopsis
zeteki has 5 denticles within the apertural lip, a posterior
knob-like denticle near the anal groove, a second large
denticle below this, followed by 3 smaller denticles. In
addition, on the anterior end of the columella are 2 con-
tiguous, elongate denticles. The first is the larger, and with
the second one, situated more deeply within the aperture,
forms a continuous ridge extending into the aperture for
a considerable distance. Anterior to this structure there is
a strong groove followed by a raised, rounded ridge at the
end of the columella.
In Muricopsis armatus the inner surface of the apertural
lip bears 5 to 7 equidistant denticles and on the columellar
edge are 2 denticles, 1 at the mouth and the second re-
cessed more deeply into the aperture.
Vol. 12; No. 3
THE VELIGER
Page 355
Figure 3
Muricopsis blainvillei (PAYRAUDEAU, 1826)
operculum, interior and exterior
In Muricopsis jaliscoensis, as noted earlier, there are,
consistently, 7 denticles on the inner surface of the aper-
tural lip; a large, knob-like one near the anal groove, at
some distance from this another denticle of moderate
strength, after another space a series of 4 closely spaced
denticles, and the final, anterior-most denticle after an-
other gap. On the columella, directly above the siphonal
canal, are 3 contiguous denticles in a posterior-anterior
orientation; the posterior one most prominent.
ACKNOWLEDGMENTS
We gratefully acknowledge our debt to the following
friends and colleagues for courtesies of various kinds in
the preparation of this paper: Dr. William K. Emerson,
Dr. James H. McLean, Mr. Lawrence E. Thomas, Mrs.
Twila Bratcher, Dr. Donald R. Shasky, Mrs. Ruth Purdy,
and Mr. and Mrs. Carl Shy. We especially wish to thank
Mr. Dallas Clites for his outstanding photographic work.
LITERATURE CITED
ApaAMSs, ARTHUR
1854. Descriptions of new shells from the collection of H.
Cuming, Esq. Proc. Zool. Soc. London for 1853 (prt.
XXI) : 69 - 74 (25 July 1854)
Apams, Henry, & ARTHUR ADAMS
1853-1858. The genera of Recent Mollusca, arranged according
to their organization. London; 1: vi-xl, 1-484; 2: 1-661: 3:
pits. 1 - 138
Bivona-BERNARDI, ANTONIO
1882. Caratteri di un nuovo genere di conchiglie della famig-
Figure 4
Muricopsis jaliscoensis Rapwin & D’ArTILIO, spec. nov.
operculum, interior and exterior
lia delle Columellarie del Signor Lamarck. Effem. Scient.
Lett. Sicilia 2: 8- 12 {not seen]
Broccui, GIovANNI BaTTISTA
1814. Conchologia fossile subapennina ... vol. 2; Milan,
Silvestro; 556 pp.
Bucqguoy, EucENE & PHILippE DAUTZENBERG
1882. in: Bucguoy, DauTZENBERG « DottFrus, Mollusques
marins du Roussillon. Paris, vol. 1. fasc. 1. Genres: Murex,
Pisania, Ranella, Triton, Cancellaria, Hadriania, Fusus, Euthria,
and Trophon, pp. 1 - 40; atlas plts. 1-5
CarPENTER, PHILIP PEARSALL
1865. | Diagnoses of new species of mollusks, from the west
tropical region of North America, principally collected by the
Rev. J. Rowell, of San Francisco. Proc. Zool. Soc. London
(for 1865) (1): 278-282 (June) [reprinted in CaRPENTER,
1872 (I): 269-275]
CossMANN, ALEXANDRE EpouARD MAURICE
1903. Essais de paléoconchyliologie comparée, vol. 5 (Paris)
(chez l’auteur): 215 pp.; 9 plts.; 16 text figs.
Dance, STANLEY PETER
1966. Shell collecting: an illustrated history.
Press, Berkeley & Los Angeles, 344 pp.; 35 plts.
Dittwyn, LEwis WESTON
1817. A descriptive catalogue of Recent shells arranged
according to the Linnaean method, with particular attention
to the synonymy. London, 1: 1 - 580; 2: 581 - 1092; index
GMELIN, JOHANN FRIEDRICH
1791. Caroli Linnaei systema naturae per regna tria naturae.
Ed. 13, aucta, reformata, Vermes Testacea Lipsiae 1 (6).
pp. 3021 - 3910
GRANGER, A.
1885. Histoire naturelle de France. pt. 6, Mollusques (Cephal-
opodes, Gasteropodes). Paris. [not seen]
HEILPRIN, ANGELO
1887. Explorations on the west coast of Florida and in the
Okeechobee wilderness. Trans. Wagner Free Inst. 1: 1 to
134; 19 plts.
Univ. Calif.
THE VELIGER
Page 356
HErTLeEIN, LEo GeorcE & ARCHIBALD McCLureE STRONG
1951. | Eastern Pacific expeditions of the New York Zoological
Society. XLIII. Mollusks from the west coast of Mexico and
Central America; PartsI-X. Zoologica, prt. 10; 36: 67 - 120;
pits. 1-11 (20 August 1951)
KEEN, A. Myra
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi + 624 pp.; illus.
Stanford Univ. Press, Stanford, Calif. (5 December 1958)
LAMARCK, JEAN-BAPTISTE PIERRE ANTOINE DE MONET DE
1822. Histoire naturelle des animaux sans vertébres, 7 [Mol-
lusques]. Paris (“‘chez l’auteur, au jardin du Roi”) pp. 1-711
(August 1822)
Locarp, ARNOULD
1892. Les coquilles marines des cétes de France, descriptions
des familles, genres et espéces. Paris; 384 pp.
MonTEROSATO, Tommaso pi Maria ALLERI
1878. | Enumerazione e sinonimia delle conchiglie Mediterranee
. pt. 1, Palermo [not seen]
Mé6rcu, Orro ANpREAs Lowson
1852-1853. Catalogus conchyliorum quae reliquit D. Alphonso
d’Aguirra et Gadea, Comes de Yoldi. Hafniae, fasc. 1: 1-170
WENZ, WILHELM
1941. Gastropoda
Vol. 12; No. 3
Patias, PETER SIMON
1774. Specilegia zoologica ..
PaYRAUDEAU, B. C.
1826. Catalogue descriptif et méthodique des annelides et
mollusques de I’Ile de Corse. 218 pp.; 8 plts. [not seen]
Risso, ANTOINE
1826. Histoire naturelle des principales productions de l’Europe
méridionale et particuliérement de celles des environs de Nice
et des Alpes maritimes; ... 4. Paris (R-G. Levrault)
pp. i- vii+1 - 439; plts. 1-12
Sowersy, GeorcE BRETTINGHAM (2%? of name)
1834. The conchological illustrations, Murex catalogue, prt.
61, p. 8; plt. 61, spec. 23
Swainson, WILLIAM
1840. A treatise on malacology; or the natural classification of
shells and shell-fish. London, i - viiit1-419; 130 text figs.
(May 1840)
., fasc. 10, Berolino [not seen]
TaTE, RALPH
1888. The gastropods of the older Tertiary of Australia.
(Part 1). Trans. and Proc. and Report, Roy. Soc. South Aus-
tralia, for 1886 - 1887, 10: 91-176; plts. i - xiii.
[not seen]
in O. H. Schindewolf (ed.): Handbuch
der Paldozoologie. Berlin, vol. 6, prt. 5: 961 - 1200
Woop, WILLIAM
1828.
Supplement to the Index Testaceologicus or a catalogue
of shells, British and foreign. London, W. Wood. 59 pp.; 8 plts.
index to plates: 34 pp.
s
Vol. 12; No. 3
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Page 357
Two New Species of Deepwater Bivalves
from the Caribbean Sea
H. E. VOKES
Department of Geology, Tulane University, New Orleans, Louisiana 70118
(Plate 53; 2 Text figures)
AMONG THE SEVERAL INTERESTING species of bivalve mol-
lusca secured by the U. S. Fish and Wildlife M/V Oregon
II during a recent program of fisheries investigations in
the Caribbean Sea were a pair of fresh valves of Solemya
(Acharax) and a complete specimen, with animal, plus a
worn right valve of a large species of Lima (Acesta). Both
proved to be undescribed species, and both represent the
second species of their respective subgenera to be recorded
from the waters of the western Atlantic Ocean. All were
secured in one shrimp net trawl from Oregon II station
10288, at latitude 11°27’ N, longitude 73°42’ W, about
20 miles off the coast of Colombia, South America, at a
depth of 220 fathoms. Judging from small amounts of
adhering matrix, the bottom sediment was composed of
greenish-grey mud.
The writer is greatly indebted to Mr. Harvey R. Bullis,
Base Director of the Pascagoula, Mississippi Station of the
Fish and Wildlife Service for permission to study and
describe these specimens.
SOLEMYIDAE Gray, 1840
Solemya Lamarck, 1818
Type species, by SD (Cuitpren, 1823) ““Solenomya”
[= Solemya] Mediterranea LaMarck = Tellina to-
gata Pout, 1791 + solen (Von Saus, 1793). Recent,
Mediterranean and Adriatic Seas.
For synonymy and a discussion of the problem of fixing
the type species of Solemya, see Vokes, 1955, which also
includes a list of known Tertiary and Recent species.
(Acharax) Dax, 1908
Type species, by OD, Solemya johnsoni Datt, 1891.
Recent, Eastern Pacific.
The subgenus Acharax Datu includes those large, usu-
ally deep water species of Solemya with a wholly external,
opisthodetic ligament.
Solemya (Acharax) caribbaea H. E. VoKEs, spec. nov.
(Plate 53, Figures 1, 2; Text figure 1)
The shell is heavy for the genus, large, elongate, strongly
inequilateral, widely gaping anteriorly, with the anterior
end much longer than the posterior and with broad, low
and inconspicuous umbones located slightly behind the
posterior third of the total length. It is covered by a strong,
blackish-brown periostracum that is produced beyond the
margins of the shell, especially anteriorly and ventrally,
and is continuous dorsally over the gap between the two
valves which do not contact each other except in the area
immediately between the umbos. This periostracum is
translucent and of a dark amber tint in the portions that
project beyond the shell; in the posterior and postero-
ventral areas it is produced into ragged finger-like proces-
ses that are prolongations of that portion of the structure
covering the wide inter-rib areas of the valve surface,
with the re-entrants between the processes making the
areas where the periostracum lies over the broad primary
ribs on the valve. In the median areas of the valve, where
the rib and interspace ornamentation is not strongly devel-
oped, the periostracal projection is not divided. The ante-
rior dorsal margin of the valve is long, slightly convex,
almost straight, sharply rounded into the anterior end, the
upper half of which is straight, the ventral half broadly
and regularly rounded into the very slightly concave vent-
ral margin which, posteriorly, curves gently into the arcu-
ate posterior margin. The posterior dorsal margin is
marked by a low narrow ridge that rises vertically above
the adjacent outer valve surface, straight in the area
behind the nymph plate, slightly arcuate laterally around
Page 358
THE VELIGER
Vol. 12; No. 3
that plate to leave a groove for the reception of the liga-
ment, then curving upward to terminate immediately
behind the umbo. The dorsal margin is the thickest area
of the valve, there being a buttress-like thickening under
the anterior half of the nymph plate that extends forward
and under the low umbone to fill the sub-umbonal cavity;
anterior to this the margin is thickened externally to form
a low, round-topped, rib-like structure that extends to the
most anterior extremity of the margin. The surface of the
valve below this dorsal ridge is ornamented by low broad
radial ribs separated by relatively wider interspaces. Three
such ribs extend to the anterior end of the valve, while a
fourth marks the antero-ventral junction. The tops of
these ribs are relatively flat in the younger stages of
growth, but in the later stages one or more (usually one)
very shallow and broad grooves tend to develop on the
rib surface; these are not as sharp nor as well marked as
are the interspaces between the ribs. These interspaces are
set off from the ribs by relatively narrow and deep round-
bottomed channels with the area between the channels
slightly convex upward, suggesting low rounded secondary
ribs and these, in turn, bear on their upper surface narrow
grooves, usually 2 or 3 in number, that serve to delimit
narrow, rounded, tertiary rib-like structures. Posterior to
the fourth primary rib the median surface of the valve
shows a tendency to have broad ribs and interspaces, but
there is no marked distinction in the relative height of
these structures though the rounded margining grooves
persist as do, to a lesser extent, the shallower, less well
marked ridges on the central parts of the rib and inter-
space area. The postero-ventral margin is marked by 3
relatively wide and flat-topped radial ribs separated from
each other by 2 broad interspaces that are approximately
of the same width as the adjacent ribs, and like the inter-
spaces on the anterior end of the valve are somewhat
convex upward in their medial areas. These 3 postero-
ventral ribs tend to project very slightly along the valve
Figure 1
Solemya (Acharax) caribbaea H. E. Voxss, spec. nov.
A: Interior of right valve of holotype. B: Same, but valve tilted
to show muscle scars that lie under the dorsal margin
aa-c anterior adductor muscle scar, “catch”segment
aa-f anterior adductor muscle scar, “fast” segment
pa-f and pa-c posterior adductor muscle scar, “fast” and “catch”
segments, respectively
apr anterior pedal retractor muscle scar
ppr posterior pedal retractor muscle scar
pe? scar of pedal elevator muscle?
(approximately X 1)
margin with the interspaces forming slight re-entrants
between them. The surface of the valve above the most
posterior of the 3 ribs is smooth, marked only by growth
lines.
Explanation of Plate 53
Solemya (Acharax) caribbaea H.E. VoxEs, spec. nov.
Figure 1: Exterior of left valve covered by the ragged, dark brown
periostracum
Figure 2: Exterior of left valve with periostracum removed (ob-
liquely lighted to emphasize radial ornamentation)
Holotype USNM 679377
Lima (Acesta) colombiana H.E. Voxes, spec. nov.
Figure 3: Anterior view of conjoined valves
Figure 4: Exterior of right valve.
Holotype USNM 679378
(all figures X 1)
THE VELIGER, Vol. 12, No. 3 [H. E. VoxEs] Plate 53
Figure 1
Figure 3 , Heures
Vol. 12; No. 3
The paravincular ligament is opisthodetic, with the
fibrous resilium seated on a relatively strong nymph plate
that rises from the inner side of the thickened margin of
the valve and curves upward so that its straight dorsal
margin is essentially parallel with the anterior dorsal
margin of the valve, though topographically a little lower
than the latter. The resilium is attached to the outer sur-
face of the nymph and arches over its dorsal side to its
attachment in the other valve. The outer lamellar tensil-
ium is longer than the resilium projecting posteriorly to
it, thick and heavy, set in a distinct groove between the
nymph plate and the raised posterior dorsal margin of
the valve, and somewhat darker in color than the peri-
ostracum to which it is firmly cemented.
The interior of the valve is chalky white in color, with
the interrib areas of the anterior and the posterio-ventral
areas weakly reflected on its surface. The weak ribbing
on the external median surface is evidenced only along
the ventral margin where the deeper grooves margining
the interspaces are represented by low rounded riblets.
Anterior adductor scar relatively large, subquadrate in
outline, not impressed; posterior adductor smaller, slight-
ly impressed. Both scars show differentiation into 2 areas,
a smaller outer and a larger inner one that probably re-
flect the areas occupied by the “quick” or “fast” muscle
segment comprised of striated fibers and those with the
“slow” or “catch” muscle segment comprised of smooth
muscle fibers. Analogy with other bivalve species sug-
gests that the smaller, outer portions of the scars were the
seat of the “slow” muscle segments. A narrow, linear
series of muscle attachment impressions follows the under
side of the thickened dorsal margin extending posteriorly
from a point immediately posterior to the anterior adduc-
tor scar to approximately the mid-length of the thickened
nymph. Small areas set off from the others by a shallow
groove, one at the anterior end of this series, the other at
the posterior end located upon the edge of the nymph,
are, by analogy with the illustrations of the anatomy of
Solemya “mediterranea LAMARCK” given by DESHAYES
(1845, plate 19, figure 5) attachment areas for the pedal
retractor muscles, while the elongate, median scar prob-
ably marks the site of the pedal elevator muscle (Text
figure 1B). Pallial line not well delimited, but appears
to have been entire, non-sinuated.
Holotype: U.S. N.M.no. 679377; length (with peri-
ostracum) 95mm; (without periostracum) 78.3 mm;
height (with periostracum) 47 mm, (without periostra-
cum) 32.3 mm; diameter (paired valves) 22 mm.
Solemya (Acharax) caribbaea, spec. nov., differs from
S. (A.) grandis VERRILL & BusH (1898, p. 885; plt. 86,
figs. 1,2), the only other west Atlantic species of Acharax,
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Page 359
described from material from 4 dredging localities off
the eastern coast of the United States between Maryland
and Massachusetts in depths of 300 to 1200 fathoms,
in being a larger species, with more posteriorly situated
umbones, and with fewer (4) radial ribs on the anterior
end, as compared with 6 to 8 on S. grandis.
Solemya (Acharax) johnsoni Datu (1891, p. 189; 1895,
p. 712; plt. 35, fig. 1) and its probable synonym (see
Woonrine, 1938, p. 27), S. (A.) agassizii (DALL, 1908, p.
365; plt. 16, fig. 10) have more numerous radial ribs
both anteriorly and on the postero-ventral area, and um-
bones that are slightly more anteriorly situated than are
those of the present species. Both are apparently larger
species. There is, however, some uncertainty as to the
dimensions of the type of S. (A.) agassizii; DALL gives the
dimensions of the species in his description as: “Length
. excluding periostracum, about 95; height 30; and
diameter, 25 mm. The species reaches a length of more
than 150 mm.” But his figured specimen (plate 16, figure
10), lacking periostracum, is said to be “lon. 145.0 mm.”
Wooprine (1938, p. 27) states that “The type of agas-
sizi is a large shell ... that has a length exclusive of the
periostracum of about 143 millimeters.” Accepting these
larger figures, the type of S. (A.) agassizii is almost twice
the size of S. (A.) caribbaea.
LimméE RaFINESQUE, 1815
Lima Brucutkre, 1797
Type species, by subsequent tautonomy (LAMaRCK,
1801) Ostrea lima Linnarus, 1758 = Lima squa-
mosa LAMARCK, 1801. Recent.
The writer has discussed in some detail the problems of the
authorship and fixation of the type species in an earlier
paper (VoKEs, 1963).
(Acesta) H. & A. Apams, 1858
Type species, by M, Lima “excavata (CHEMNITZ)” =
Ostrea excavata Fasricius. Recent, 150 to 1450 fath-
oms, Greenland to the Canary Islands.
A few years ago the writer described (VoKEs, 1963, p.
77) under the name Lima (Acesta) bullisi a large species
that had been trawled by the Fish and Wildlife Service
M/V Oregon from localities 70 miles south-southeast, and
75 miles due south of the entrance to Mobile Bay, Ala-
bama, at depths ranging from 300 to 600 fathoms. Subse-
quently he has seen, in the private collection of Mrs.
Page 360
Mildred Tate of Lake Jackson, Texas, a very fine pair of
valves that had been trawled by a shrimp boat from below
226 fathoms off the Mississippi River delta. This species,
the first recorded from western Atlantic waters, has,
to date, been recorded only from the fauna of the northern
part of the Gulf of Mexico.
Lima (Acesta) colombiana H. E. VokEs, spec. nov.
(Plate 50, Figures 3, 4; Text figure 2)
Shell large, elongate ovate in outline, very inequilateral,
almost equivalved, with the right valve slightly more in-
flated than the left; umbones low, almost terminal ante-
riorly, but slightly prosogyrate; anterior ear small, poste-
rior elongate, not delimited from rest of valve; hinge line
straight, relatively short (approximately 31 mm in length),
with a strongly curved ligamental pit that is 11.8mm
wide where it intersects the hinge line; elongate lunular
area excavate, its outer portion marked by fine, irregular
radial ribs, its inner side strongly corrugated by growth
lamellae; the anterior margin is broadly concave in the
lunular area, but the anterior-dorsal slope of the valve,
including the area above the lunule, is almost straight,
anterior end rather sharply rounded, ventral and posterior
ends broadly and regularly rounded, posterior dorsal slope
Figure 2
Outline of right valve of Lima (Acesta) colombiana H. E. VoxEs,
spec. nov. (continuous line) compared with that of Lima (Acesta)
bullist H. E. Voxes, 1963 (broken line). Both approximately
one-half natural size
THE VELIGER
Vol. 12; No. 3
slightly convex, almost straight, subangulate at the junc-
tion with the short, relatively straight dorsal margin.
Surface ornamentation characteristic of the subgenus,
with, on the median surface of the valve, numerous fine,
irregular, round-topped radial riblets of variable width
separated by exceedingly narrow irregular grooves; the
ribs on the dorsal area so fine as to be visible only under
oblique light and magnification, increasing in relative
width and strength as the shell becomes larger; number of
ribs increases by development of median groove on the
rib surface which later assumes the strength of a narrow
interspace; laterally the ribs become relatively narrower
with progressively wider, rounded interspaces, until near
the margin they are best described as low, rounded radial
threads with interspaces 3 to 4 times as wide as the ribs.
Interior of valve with smooth margins, adductor scar
situated relatively high in the valve, rounded to sub-tri-
gonal in outline, not impressed, its anterior margin at
the mid-width of the valve.
Holotype: U.S. N. M. no. 679378, height 124 mm, length
94.8 mm, diameter (paired valves) 60.3 mm.
Paratype: U.S.N,M. no. 679378a, height 120mm,
length (incomplete) 91.4mm, diameter (right valve)
32.6 mm.
Lima (Acesta) colombiana spec. nov. differs from L.
(A.) bullisi Voxes in having a shell that is much higher
in proportion to its length (see Text figure 2), with a
longer and straighter antero-dorsal margin and more
sharply rounded anterior end. The surface ornamentation,
while characteristic of the subgenus in both forms, is
more weakly developed and finer in the new species than
in the Gulf of Mexico form, there being 13 to 14 ribs per
centimeter on the median area of the valve as compared
with 8 to 9 per centimeter in L. (A.) bullist.
The Oligocene Lima (Acesta) goajira Otsson & RicH-
Arps (1961, p. 6; plt. 1, fig. 1), described from the adja-
cent Goajira Peninsula of Colombia, judging from the
growth lines, appears to have had a somewhat comparable
outline, although the anterior end may have been more
broadly and regularly rounded. This form is described,
however, as being smooth and devoid of radial omamenta-
tion except for a “narrow area of wavy, radial threads
along the posterior dorsal margin.”
Of the Recent Eastern Pacific species, Lima (Acesta)
agassizi DauL, 1902, dredged from 322 fathoms in the
Gulf of Panama, has, judging from the illustrations (DALL,
1908, p. 47; plt. 16, fig. 1), a shorter antero-dorsal margin,
longer and more broadly rounded anterior end, and
coarser radial ornamentation; L. (A.) mori HERTLEIN
(1952, p. 379; plt. 20, figs. 12, 13), described from Mul-
berry Seamount, off San Mateo County, California, is
Vol. 12; No. 3
much more rounded in outline, with very low and ap-
parently inflated umbone, and so far as can be judged
from the illustration, coarser radial ornamentation.
The recently described mid-Pacific Acesta virgo HaBE
& OxuTant (1968, p. 51; plt. 3, fig. 8) from 480 meters
off Midway Island, has an outline that is quite like that
of L. (A.) colombiana with a similarly long and relatively
straight antero-dorsal slope and broadly rounded ventral
and posterior margins. It differs, however, in its shorter,
more obliquely sloping postero-dorsal and shorter dorsal
margins. The ornamentation, not well shown in the illus-
tration, is described as consisting of regularly spaced
radial riblets, a condition that would differ greatly from
the irregularity of the ribbing on the present species.
LITERATURE CITED
Dati, WILLIAM HEALEY
1891. Scientific results of explorations by the U.S. Fish Com-
mission steamer Albatross. XX. — On some new or interesting
West American shells obtained from the dredgings of the U.S.
Fish Commission steamer Albatross in 1888 and from other
sources. Proc. U.S. Nat. Mus. 14 (849): 173-191; plts.
5-7 (24 July 1891)
1908. Reports on the dredging operations off the west coast
of Central America to the Galapagos, to the west coast of
Mexico, and in the Gulf of California. . XIV. The
Mollusca and Brachiopoda.
Harvard 43 (6): 205 - 487; 22 plts.
Bull. Mus. Comp. Zool.,
(October 1908)
THE VELIGER
Page 361
DesHAyYEs, GzrRaRD PAuL
1844 [1845] - 1849. Explorations scientifiques de l’Algérie pendant
. 1840-1842. ... Zoologie. I. Histoire naturelle des Mol-
lusques. Atlas: 160 pp.; 155 col. plts. Paris, France
Hase, TADASHIGE & TAKASHI OKUTANI
1968. Some new and interesting shells from the sea around
Midway Island. Venus, Japan. Journ. Malacol. 27 (2) :
47 - 56; pit. 3 (October 1968)
HERTLEIN, LEO GEORGE
1952. Description of a new pelecypod of the genus Lima from
deep water off Central California. Proc. Calif. Acad. Sci.
(4) 27 (12): 377 - 381; plt. 20 (11 July 1952)
Oxsson, AxEL ADOLF & Horace GARDINER RICHARDS
1961. Some Tertiary fossils from the Goajira Peninsula, Co-
lombia. Notulae Naturae no. 350: 1-16; 2 plts.
(10 November 1961)
VeRRILL, AppISON E. &« KATHARINE J. BusH
1898. Revision of the deep-water Mollusca of the Atlantic
Coast of North America, with descriptions of new genera and
species. Part I. — Bivalvia. Proc. U.S. Nat. Mus. 20:
775 - 901; plts. 71 - 97 (15 June 1898)
VoKEs, Haro_p ERNEST
1955. Notes on Tertiary and Recent Solemyacidae.
Paleont. 29 (3): 534 - 545
1963. Studies on Tertiary and Recent giant Limidae. I. A new
species of Acesta from the Gulf of Mexico. II. A catalogue of
the described and Recent and Tertiary species of Acesta. III.
Plicacesta, a new subgenus of Lima. Tulane Stud. Geol.
1 (2): 73-92; 2 plts. (18 January 1963)
Wooprinc, WENDELL PHILLIPS
1938. | Lower Pliocene mollusks and echinoids from the Los
Angeles Basin, California. U.S. Geol. Survey, Prof. Paper
190: i-ii+ 1-67; 9 plts.; 2 figs.
Journ.
Page 362
THE VELIGER
Vol. 12; No. 3
Descriptions of a New Genus and Eight New Species
of Eastern Pacific Fissurellidae, with Notes on Other Species
JAMES H. McLEAN
Los Angeles County Museum of Natural History
goo Exposition Boulevard, Los Angeles, California 90007
(Plate 54; 1 Text figure)
THE EASTERN PACIFIC FISSURELLDAE are known chiefly
through Pirssry’s monograph in the Manual of Con-
chology (1890). More recently the species of the Panamic
province were treated by Keen (1958) and those of the
northeastern Pacific in a dissertation by McLEan (1966).
A comprehensive review of the entire family similar to
that for the western Atlantic by FARFANTE (1943a, 1943b,
1947) is not available. Most of the genera discussed by
FaRFANTE are represented in the eastern Pacific and these
papers are of considerable value for comparison.
In assisting with the treatment of Fissurellidae for the
forthcoming revised edition of “Seashells of Tropical West
America,” by Dr. Myra Keen, I have realized that there
is a need for the description of a new genus and several
new species. These descriptions are presented here.
Five of the new species are from the tropical Panamic
faunal province, while 3 are from the south temperate
Peruvian province. Two of the Panamic species are from
the Galapagos Islands, approximately 600 miles west of
Ecuador. The 3 species from the Peruvian province are
from the offshore Chilean island groups Juan Fernandez
and San Felix. These islands bear much the same relation-
ship to the Peruvian province as do the Galapagos Islands
to the Panamic province. Many of the species are en-
demic. ODHNER (1922) listed 39 species from shallow
water at the Juan Fernandez Islands, 50% of which were
endemic. Very little is known of the offshore mollusks of
the Chilean islands. The 3 species described herein result
from collecting on the Anton Bruun Cruise 17 that visited
these islands in July 1966.
Institutions mentioned in the text and their abbrevia-
tions are as follows:
AHF - Allan Hancock Foundation (gastropod collec-
tion on loan to LACM)
CAS - California Academy of Sciences, San Francisco
LACM -— Los Angeles County Museum of Natural
History
SU — Stanford University, Stanford, California
USNM - United States National Museum,
Washington, D. C.
Tugalt chilensis McLean, spec. nov.
(Plate 54, Figures 8, 9)
Description of Holotype: Shell small, elongate ovate,
with nearly parallel sides, moderately elevated. Apex
blunt, nucleus worn smooth, directed posteriorly, + the
length of the shell from the posterior margin. Anterior
slope convex, posterior slope concave, sides nearly flat.
On a level surface the sides of the shell are slightly raised
relative to the ends. Sculpture consists of radial and con-
centric ribbing of nearly equal strength, beaded at inter-
sections and producing indistinct square cancellations.
Concentric growth irregularities are also apparent. Ap-
proximately 30 primary ribs diverge from the apex and
secondary ribs appear at half its final size, resulting in
about 60 ribs of identical size and strength that wrap
around the thick margin of the shell. No selenizone or
crenulation apparent at the anterior margin. Interior of
holotype encrusted, characters of muscle scar not discern-
ible. Length, 10.4mm, width 6.5mm, height 3.8mm
(holotype) .
Type Material: Holotype, LACM 1306.
Type Locality: Carvajal Bay, south end of Isla Mas a
Tierra, Juan Fernandez Islands, Chile, 34°50’S, 79°00’ W.
A single dead specimen was collected at low tide by Mr.
Roger Seapy of the University of Southern California, 17
July 1966, Anton Bruun, Cruise 17.
Discussion: Although represented by but a single shell in
poor condition, it is adequate for subsequent recognition
and its description emphasizes the presence of this Austral-
asian genus in the Eastern Pacific fauna. Tugali chilensis
Vol. 12; No. 3
is a species closely related to T: suteri THEE, 1916, of
the New Zealand fauna. Specimens of the latter species on
hand are slightly larger, have more numerous ribs and
the apex at 3 the distance from.the posterior margin.
The Juan Fernandez Islands are approximately 400
miles west of the mainland of Chile at the latitude of
Valparaiso.
Nesta galapagensis McLean, spec. nov.
(Plate 54, Figures 18, 19)
Description of Holotype: Shell small, white, base elongate
ovate. Nucleus extremely minute, at the posterior end of
the shell just above the margin, twisted to the right (in
apertural view). The anterior face of the shell forms the
entire dorsal surface, the posterior face represented by a
narrow shelf, as in the hinge area of a bivalve. A fairly
broad selenizone extends from the nucleus to the anterior
margin, bounded by raised ridges and provided with U-
shaped incremental lines. Sculpture consists of numerous
radial and concentric ribs of equal strength, forming
squarish cancellations. The radial ribs on crossing the
concentric ribs are drawn out in blunt lamellar points,
visible only in lighting from the direction of the apex.
Entire anterior margin broken in holotype. Interior glossy,
with heavy callus deposition along the line of the seleni-
zone. Length, 5.5mm, width, 3.3mm, height 1.7mm
(holotype).
Type Material: Holotype, LACM - AHF 1307.
Type Locality: Tagus Cove, Albemarle (Isabella) Island,
Galapagos Islands, Ecuador, 0°16’S, 91°22’30” W, 80-100
fathoms, Velero III bottom sample station 432, 15 Janu-
ary 1934, 1 specimen.
Discussion: Nesta galapagensis is the first reported spe-
cies of Nesta H. Apams, 1870, in the Eastern Pacific.
According to FARFANTE (1947, p. 98) there are two other
species of Nesta, the strongly sculptured type species, N.
candida H. Apams, 1870, from the Red Sea, and the
nearly smooth sculptured N. atlantica FARFANTE, 1947,
from Florida. Having concentric sculpture, N. galapag-
ensis is therefore closer to the type species, but the apex
in that species curves below the posterior margin.
Emarginula Lamarck, 1801
The genus Emarginula was unknown in the Eastern Pacif-
ic until the description of E. velascoensis SHasxy, 1961,
from the Gulf of California (SHasxy, 1961, p. 18).
THE VELIGER
Page 363
A second tropical west American species represented by
4 specimens from the Galapagos Islands and Colombia
taken at depths of 45 to 100 fathoms answers the descrip-
tion of E. tuberculosa Ligassi, 1859, given by FARFANTE
(1947, p. 100; plt. 44, figs. 1-7). This is a relatively
large species, the largest specimen from 45 fathoms, off
Octavia Rocks, Colombia, measures 13.4 mm in length
and 7.2 mm in height (Plate 54, Figures 10, 11). Sculp-
ture consists of numerous fine radial and concentric ribs
producing beaded intersections and deep square pits. Every
4" rib at the margin, each of which represents an original
primary rib diverging from the apex, is slightly more
prominent than adjacent secondary ribs, as in FARFANTE’S
illustrations. No essential points of difference can be de-
tected. Emarginula tuberculosa is also represented in the
eastern Atlantic off Portugal and the Azores and its type
locality is the Sicilian Miocene. It is therefore a species
of considerable antiquity and broad distribution else-
where; its presence in the Eastern Pacific is not unex-
pected.
Descriptions of two additional eastern Pacific species
follow. These two species were taken in the same dredge
haul at a depth of about 90 fathoms at San Felix Island,
approximately 550 miles off the coast of central Chile,
and therefore are members of the south temperate or
Peruvian faunal province.
An anatomical observation pertaining to at least two
species of Emarginula may be made here. Cowan (1969)
observed that in Fissurisepta pacifica Cowan, 1969, the
ctenidium is monopectinate and attached to the roof of
the mantle cavity, a condition not previously reported in
Fissurellidae. One specimen of E. tuberculosa from Tagus
Cove, Isabella Island, Galapagos has a dried animal and
the holotype of E. angusta, new species, also has the
animal dried in place. Both specimens have monopectin-
ate ctenidia resembling those illustrated by Cowan,
suggesting that this structure of the ctenidium may be
more widespread in the emarginuline fissurellids than has
been supposed.
Emarginula angusta McLEAN, spec. nov.
(Plate 54, Figures 14, 15)
Description of Holotype: Shell small, sturdy, yellowish
white, base elongate-oval, with nearly parallel sides;
height about 3/5 the length, apex posterior, directly above
the posterior margin of the shell. Anterior slope broadly
convex, posterior slope convex below the apex, sides of
shell nearly straight. Nucleus minute, turned under the
succeeding part of the shell. Sculpture consists of 16
strong primary cords diverging from the apex; secondary
Page 364
THE VELIGER
Vol. 12; No. 3
cords develop on the anterior and lateral slopes and reach
nearly the strength of the primary cords at the margin;
concentric sculpture consists of raised ridges that reach
only half the thickness of the primary ribs but form beads
at intersections and deep squarish pits. Margin crenulated
by rib extensions. The slit is long and narrow, about 1/5
the length of the anterior slope; the lamellae along the
fasciole are thickened, equal in number and thickness to
the concentric ridges. Interior glossy, dried animal re-
maining in the shell. Length, 6.8mm, width, 4.6 mm,
height, 4.1 mm (holotype).
Type Material: Holotype, LACM 1308.
Type Locality: Off San Felix Island, Chile, 26°20’ S,
80°02’ W, 170-160 meters (93-87 fathoms), Anton
Bruun, Cruise 17 station 676B, Menzies trawl, 12 July,
1966, 1 specimen.
Discussion: Emarginula angusta differs from other East-
ern Pacific and Western Atlantic species in having an
elongate and narrow basal outline, hence the name, mean-
ing narrow. Other species have a more oval basal outline.
In addition, the lamellae of the fasciole in E. angusta
are broad and thick, unlike the other species that have
thin lamellae.
Emarginula dictya McLEan, spec. nov.
(Plate 54, Figures 16, 17)
Description of Holotype: Shell small, fragile, white, base
ovate, height about 4 the length, apex posterior, directly
above the posterior margin of the shell. Anterior slope
convex, posterior slope flat sided below the apex. Nuc-
leus minute, greatly overhung by the succeeding part of
the shell. Sculpture consists of 20 primary cords diverg-
ing from the apex; secondary cords develop on the
anterior and lateral faces but do not reach the strength
of the primary cords at the margin; concentric sculp-
ture consists of narrow raised ridges that do not cross the
primary ribs. Margin crenulated by rib extensions. The
slit is long and narrow, almost 4 the length of the anterior
slope; the lamellae along the fasciole are thin, raised
nearly to the margins of the fasciole. Interior glossy,
thickened near the slit, anal fasciole marked by thickened
callus. Length, 4.6mm, width, 3.6mm, height, 2.3 mm
(holotype).
Type Material: Holotype, LACM 1309; 1 paratype LA
CM 1310.
Type Locality: Off San Felix Island, Chile, 26°20’S,
80°02’ W, 170-160 meters (93-87 fathoms), Anton
Bruun, Cruise 17 station 676B, Menzies trawl, 12 July,
1966, 2 dead and slightly broken specimens.
Discussion: Emarginula dictya differs from the 3 other
eastern Pacific species of Emarginula in having weak
concentric ribbing and consequently lacking the beaded
sculpture at intersections of the ribs. It resembles E. sicula
Gray, 1825, of the Western Atlantic and Mediterranean.
Emarginula sicula, however, is larger, more elevated, and
the apex is near the summit rather than below the
mid-height position as in E. dictya.
The name is derived from the Greek noun, dictyon,
net, with reference to the fine reticulate sculpture.
Rimula DrEFrRance, 1827
The genus Rimula was established in the Eastern Pacific
with the description of R. californiana Berry, 1964 from
Catalina Island. As this species has not been illustrated,
the holotype is figured here (Plate 54, Figure 3). Addi-
tional specimens are known from Guadalupe Island (LA
CM 65-42) and San Martin Island, Baja California
(CAS 24041), at depths of 10-20 fathoms on gravel
bottoms.
KEEN (1968, p. 403) showed that Rimula mazatlanica
CaRPENTER, 1857, represents a juvenile Diodora. A
second eastern Pacific Rimula may now be added.
Explanation of Plate 54
Figures 1, 2: Fissurella (Cremides) decemcostata McLean, spec.
nov. Holotype LACM no. 1313. Acapulco, Mexico XK 14
Figure 3: Rimula californiana Berry, 1964. Holotype, SU 9500.
Catalina Island, California X6
Figure 4: Rimula astricta McLEan, spec. nov. Holotype USNM
no. 267019. San Esteban Island, Mexico x6
Figures 5, 6, 7: Diodora punctifissa McLean, spec. nov. Holotype
LACM - AHF no. 1311. Wenman Island, Galapagos Islands X 4
Figures 8, 9: Tugali chilensis McLzan, spec. nov. Holotype LACM
no. 1306. Juan Fernandez Islands, Chile. x4
Figures 10, 11: Emarginula tuberculosa Lrpassi, 1859. AHF no.
431-35. Octavia Rocks, Colombia X2
Figures 12, 13: Leurolepas roseola McLEan, spec. nov. Holotype
LACM - AHF no. 1315. Tres Marias Islands, Mexico X 3
Figures 14, 15: Emarginula angusta McLEan, spec. nov. Holotype
LACM no. 1308. San Felix Island, Chile x5
Figures 16, 17: Emarginula dictya McLean, spec. nov. Holotype
LACM no. 1309. San Felix Island, Chile X5
Figures 18, 19: Nesta galapagensis McLEan, spec. nov. Holotype
LACM no. 1307. Isabella Island, Galapagos Islands x5
Tue VELIGER, Vol. 12, No. 3 [McLean] Plate 54
Figure 2
Figure 13 Figure 14 Figure 15
Figure 16 Figure 17 Figure 18 Figure 19
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a oa =
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if foe. SES koe
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- = e . mas
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i x ¥
Baw <
cr ii 3
ae i.
-
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Vol. 12; No. 3
Rimula astricta McLean, spec. nov.
(Plate 54, Figure 4)
Description of Holotype: Shell small, fragile, white, base
elongate-ovate, with nearly parallel sides; height less than
4 the length. Anterior slope convex, posterior slope con-
cave. Nucleus of one small whorl, placed 1/6 the length
of the shell from the posterior margin. Fissure small, its
squared off posterior extremity at mid position on the
anterior slope, its anterior extremity drawn out into a
long tapered slit. Fasciole extending from apex to fissure,
provided with lamellae, fissure and fasciole bordered by
raised ridges which coalesce anterior to the fissure. Sculp-
ture consists of approximately 32 primary ribs, crossed by
concentric ridges of nearly equal strength, weakly beaded
at intersections and forming deep squarish pits; a few
secondary ribs arise but do not reach the strength of the
primary ribs at the margin. Margin finely crenulated by
rib extensions. Interior lustrous, translucent, the concent-
ric sculpture and fasciole of the outer surface visible from
within, Length. 5.3 mm, width, 3.3 mm, height, 1.9 mm
(holotype).
Type Material: Holotype, USNM 267019.
Type Locality: East side of San Esteban Island, Gulf of
California, 28°41’ N; 112°34’ W. The single specimen was
collected by Paul Bartsch, 13-14 April, 1911.
Referred Material: LeRoy Poorman Collection, Pasa-
dena, California, 1 specimen dredged at 17 fathoms off
San Carlos Bay, Guaymas, Sonora, April, 1965. This spe-
cimen has a large hole in the shell but the margin is
intact. It measures: length, 6.4mm, width, 3.9mm,
height, 2.0 mm.
Discussion: Rimula astricta differs from R. californiana
in having a sharp rather than blunt anterior termination
of the fissure. This is the most distinctive feature of R.
astricta, upon which the name, meaning drawn together,
tight, or narrow, is based. Rimula astricta also has a
more elongate basal outline. The only elongate western
Atlantic species is R. frenulata Dati, 1889, but this spe-
cies does not have the pointed fissure nor the coalesced
ribs anterior to the fissure.
Diodora punctifissa McLEaN, spec. nov.
(Plate 54, Figures 5 to 7)
Description of Holotype: Shell small, sturdy, white, base
oval, sides of shell raised relative to the ends, height
about 2 of the length. Apex intact, blunt pointed, erect,
THE VELIGER
Page 365
about 4 the length of the shell from the front margin.
Fissure small, just anterior to the apex, anterior end of
fissure rounded, posterior end broader. Sculpture consists
of about 30 primary ribs originating on the apex; second-
ary ribs emerge close to the apex and assume nearly
equal strength at the margin. Concentric sculpture of
weak ridges, somewhat irregular ; cancellate pitting strong-
ly developed only on the early growth stage of the shell.
Interior callus broad, extending more than + the length
of the shell, markedly truncate posteriorly. Length, 9.5
mm, width, 7.0 mm, height, 6.0 mm (holotype).
Type Material: LACM -—AHF 1311; 5 paratypes, LA
CM —- AHF 1312; 1 paratype, CAS 13276; 1 paratype,
USNM 679561.
Type Locality: Off Wenman Island, Galapagos Islands,
Ecuador, 1°23’10”N, 91°48’45” W, 100-150 fathoms,
Velero III station 143-34, 11 January 1934, 8 dead speci-
mens. The paratype specimens are immature, the largest
measures 7.6mm in length. The species may live at a
lesser depth.
Discussion: Diodora punctifissa differs from all species
of Diodora known to me in having a fully intact apex,
not absorbed by the fissure. The fissure is extremely small
and is located on the anterior slope of the shell as in the
genus Puncturella. The posteriorly truncate internal cal-
lus possessed by D. punctifissa is the main diagnostic
feature of Diodora. Other species of Diodora having
partially intact apices are known.
The name meaning small cleft calls attention to the
smallness of the fissure.
Fissurella (Cremides) decemcostata McLEAN, spec. nov.
(Plate 54, Figures 1, 2)
Description of Holotype: Shell of medium size, sturdy,
depressed. Fissure just forward of center, oblong, faintly
tripartite, about 1/9 the length of the shell. Sculpture of
10 broad, raised, nodular ribs, the 2 anterior ribs closely
adjacent. The ribs project at the margin, lines connecting
their terminations are slightly concave. Fine irregular
striae are present on the rib surfaces and interspaces.
Concentric sculpture limited to growth irregularities, re-
sulting in broad nodules on the ribs. Color whitish, the
channels separating the fine riblets dark brown where the
surface is not worn smooth. Interior glossy, greenish white,
callus area blue gray, bordered with reddish brown, inte-
rior with reddish brown stains extending anteriorly and
posteriorly from the callus to the edge of the muscle im-
pression scar. Length, 28.2 mm, width, 18.5 mm, height,
5.8mm (holotype).
Page 366
Type Material: Holotype, LACM 1313; 1 paratype, LA
CM 1314; 1 paratype, USNM 679562.
Type Locality: Playa Caleta, Acapulco, Guerrero, Mexi-
co, 16°51’ N, 99°55’ W. The type lot consisting of 3 speci-
mens was collected by the late Earl C. Huffman, 18
December, 1937.
Referred Material: Numerous lots are in museum collec-
tions from the Tres Marias Islands, Mexico (LACM
65-13), to Oaxaca, Mexico. The species occurs on flat,
exposed reefs in the intertidal zone. The upper surface is
frequently abraded and usually covered with dense tufts
of red algae.
Discussion: Fissurella decemcostata is the only eastern
Pacific species having primary sculpture of 10 broad ribs.
Fissurella microtrema SowErRByY, 1835, is a closely related
species sharing the same habitat but this species typically
has 3 strong anterior ribs and irregular strong ribbing
throughout. The only other species having the irregular
purple staining in the interior is F rubropicta Pitssry,
1890, but it differs in having a more elevated shell with
strong ribbing throughout. The irregular purple staining
of F decemcostata is invariably present and immediately
serves to separate shells from F microtrema.
Leurolepas McLean, gen. nov.
Type Species: Leurolepas roseola McLEan, spec. nov.
Diagnosis: Shell small, moderately elevated, basal outline
oblong, sides nearly parallel; margin in one plane, aper-
ture large, oval, just anterior to center, radial sculpture
lacking, concentric sculpture of thin, raised lamellae; col-
or reddish brown with darker rays; margin thick, rounded,
smooth; apertural callus narrow, encircled by a distinct
suture; muscle attachment area close to internal margin.
Animal not capable of retracting within the shell, 14
times the length of the shell (preserved), mantle envelop-
ing the shell margin on all sides, covering } the area of
the shell, extending down to cover the head and foot,
surface of mantle with minute pustules, pigmentation
pattern matching that of the shell; snout 4 the length of
the tentacles, sides of foot weakly pigmented, with an
epipodial row of tongue-shaped papillae.
Radula fissurelline, rachidian tooth similar in size and
shape to the adjacent lateral teeth, outermost lateral
large, bicuspid.
Discussion: The small rachidian tooth of Leurolepas
’ serves to place it in the subfamily Fissurellinae as delim-
ited by THIELE, 1929, in company only with the genus
THE VELIGER
Vol. 12; No. 3
Fissurella BrucuttrE, 1789, and such Australasian genera
as Amblychilepas Pitspry, 1890, and Macrochisma G. B.
Sowersy, 1839, in which the animals are also not capable
of retraction within the shell. The other eastern Pacific
and western Atlantic genera with large animals such as
Fissurellidea Orsicny, 1841, Lucapina G. B. SowERsy,
1835, Lucapinella Prtspry, 1890, Megathura Pissry,
1890, and Megatebennus Puspry, 1890, all have an
extremely broad rachidian tooth and are thereby relegated
to the subfamily Fissurellidinae as delimited by McLEAN
(1966).
The presence of a member of this otherwise Australasi-
an group of genera in the Eastern Pacific is noteworthy.
Leurolepas differs from Amblychilepas and Macrochisma
and subgenera thereof treated by Moore (1960) in having
the margin of the shell not greatly thickened internally
and in having a margin resting entirely in one plane,
rather than raised at both ends. On shell characters it
differs from all other eastern Pacific fissurellid genera in
lacking radial ribbing.
The name is derived from the Greek words leuros,
smooth, and lepas, limpet, suggested by the lack of radial
sculpture.
Leurolepas roseola McLean, spec. nov.
(Plate 54, Figures 12, 13)
Description of Holotype: Shell small, moderately ele-
vated, thin but sturdy. Anterior and lateral slopes nearly
straight sided, posterior slope convex. Fissure large, + the
length of the shell, forward of center, its posterior end at
approximately the midpoint, sides of fissure not raised.
Radial sculpture lacking except for minute striae visible
under magnification; concentric sculpture of thin, raised
growth lamellae. Surface translucent pink, with fine white
flecking and broad lateral bands of tan. Margin smooth,
rounded, slightly thickened but not offset on the inner
side, a narrow muscle impression area borders the margin
on the lateral and posterior sides. Internal callus around
fissure narrow, rounded, separated from the interior by
a distinct suture. Length, 11.2 mm, width, 6.7 mm, height,
2.7mm (holotype). :
Radula (Text figure 1): Rachidian and 3 adjacent lat-
erals of nearly the same size with simple cutting edges,
4* lateral lacking cutting edge, 5™ lateral large, bicuspid,
marginal teeth numerous, finely serrate at tips (AHF
132-34).
Type Material: Holotype, LACM-AHF 1315 (intact
animal preserved separately).
Vol. 12; No. 3
THE VELIGER
Page 367
Figure 1
Radula of Leurolepas roseola McLEan, spec. nov.
Type Locality: Magdalena Island, Tres Marias Islands,
Mexico, 21°25’40” N, 106°21’10” W, 13 fathoms, Velero
III station 970-39, 9 May, 1939, 1 specimen.
Referred Material: SU 50755, Espiritu Santo Island,
Gulf of California, 14-25 fathoms, 3 specimens; CAS
27525, Clarion Island, Revillagigedo Islands, Mexico,
1 specimen; LACM A375, Clarion Island, 20 - 40 fath-
oms, 1 specimen; AHF 129-34, Socorro Island, Revilla-
gigedo Islands, 14-18 fathoms, 3 specimens; AHF
132-34, Socorro Island, 40 fathoms, 1 specimen; US
NM _ 567722, Mazatlan, 1 specimen; LACM 65-15,
Banderas Bay, Mexico, 1 specimen; LACM 68-41,
Barra de Navidad, Jalisco, Mexico, 1 preserved specimen
diving at 30 feet; CAS 17832, Port Guatulco, Mexico, 5
specimens; LACM 65-21, Isla Otoque, Panama Bay,
5 - 15 fathoms, 1 specimen; AHF 213-34, La Plata, Ecua-
dor, 7 - 10 fathoms, 3 specimens; AHF 221-34, Gorgona
Island, Colombia, 20 fathoms, 1 specimen.
Discussion: Leurolepas roseola is evidently not uncom-
mon throughout the Panamic province on rocky bottoms
in relatively shallow sublittoral depths. It is the only
eastern Pacific fissurellid lacking radial ribbing and is
easily recognized by its large foramen and pink coloration,
which its name emphasizes.
LITERATURE CITED
BERRY, SAMUEL STILLMAN
1964. Notices of new eastern Pacific Mollusca. — VI.
Leafl. Malacol. 1 (24): 147 - 154 (29 July 1964)
Cowan, Ian McTaccartT
1969. A new species of gastropod (Fissurellidae, Fissurisepta)
from the eastern North Pacific Ocean. The Veliger 12 (1) :
24 - 26; 3 text figs. (1 July 1969)
FarFANTE, ISABEL PEREZ
1943a. The genera Fissurella, Lucapina, and Lucapinella in the
Western Atlantic. Johnsonia 1 (10): 1-20; plts. 1-5
(7 August 1943)
1943b. The genus Diodora in the Western Atlantic. John-
sonia 1 (11): 1-20; plts. 1-6 (29 December 1943)
1947. The genera Zeidora, Nesta, Emarginula, Rimula, and
Puncturella in the Western Atlantic. Johnsonia 2 (24):
93 - 148; plts 41 - 64 (7 July 1947)
KEEN, A. Myra
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi + 624 pp.; illus,
Stanford, Calif. (Stanford Univ. Press)
1968. | West American mollusk types at the British Museum
(Natural History). - IV. Carpenters Mazatlan collection.
The Veliger 10 (4): 389-439; plts. 55-59; 171 text figs.
(1 April 1968)
McLzan, James HamILTON
1966. | West American prosobranch Gastropoda: superfamilies
Patellacea, Pleurotomariacea, and Fissurellacea. PhD!
thesis, Biology, Stanford Univ., Stanford, Calif.
pp. i-x+1- 255; plts. 1-7
Moore, RaymMonp Cecit (editor and director)
1960. ‘Treatise on invertebrate paleontology. Part I (Mollus-
ca 1): xiii + 351 pp.; figs. 1 - 216
Opune_er, Nits HyaLmar
1922. Mollusca from Juan Fernandez and Easter Island. pp.
219 - 254; plts. 8,9 In The natural history of Juan Fernan-
dez and Easter Island, ed. Carl Skottsberg, vol. 3. Uppsala
Pitspry, Henry Aucustus
1890. Manual of conchology. 12: v-xii, 1-323 ;plts. 1-65.
Suasky, Donatp R.
1961. | New deep water mollusks from the Gulf of California.
The Veliger 4 (1): 18-21; plt. 4, figs. 1-10 (1 July 1961)
(June 1966)
Page 368
THE VELIGER
Vol. 12; No. 3
On Crrnouorsky’s Designation of a Lectotype
- for Murex mancinella LINNAEUS
BY
H. E. VOKES
Department of Geology, Tulane University, New Orleans, Louisiana 70118
Member, International Commission on Zoological Nomenclature
THE WRITER WAS MUCH DISTURBED to note that Dr. W. O.
Cernohorsky (1969, p. 297) had designated as lectotype
of Murex mancinella Linnaeus, 1758 a specimen in the
Linnaean collection of the Linnean Society of London
which, on the basis of all evidence is not part of the
original type material and hence is not available for
such designation. Under the provisions of the Code of
Zoological Nomenclature [Article 74(a) (i) ] CERNOHORS-
Ky’s designation is, accordingly, invalid. This, added to
the fact that the problem of the identity of the type species
of the genus Mancinella Linx, 1807, is presently being
considered by the International Commission on Zoologic-
al Nomenclature, makes his action peculiarly unfortunate
at this time.
It has long been known that the Linnaean collection
contained three specimens identified as this species. Ac-
cording to SmirH (1913, p. 287) “They were mounted
upon wooden tablets by Mr. Hanley when he wrote his
work I psa Linnaei Conchylia, and two of them are marked
in Linné’s handwriting [emphasis mine] with the number
544 of the twelfth edition of the Systema Naturae.”’ These
two are the specimens of Drupa cornus R6pine, 1798 that
CERNOHORSKY States: “‘are excluded [apparently by him]
from the type series; they were either added by Linnaeus
after 1758 or were erroneously selected by Hanley as
types of Murex mancinella.” It should be noted in passing
that Hanley did not select these specimens as types of the
species, he simply stated that they were “marked” by
Linnaeus as examples of M. mancinella, although it is
clear from his introduction to his work (cited later in the
present paper) that he considered those specimens marked
with the numbers of the tenth edition as representing the
Linnaean types of the species described in that edition.
It is to be noted, however, that DopcE (1957, p. 136)
states that these two specimens are to be taken as syn-
types of Murex mancinella, saying: ““The systematists who
- follow the current practice of retaining well-known names
whenever possible must, in the present case, base their
opinion on the presence in the collection of a single undoc-
umented specimen of the mancinella of Lamarck and au-
thors and thus retain the name mancinella, selecting that
specimen out of the syntypic lot as the lectotype of M.
mancinella Linné. This is not only a violation of the Rule
of Priority, but is the choice of a type unsupported by any
evidence whatsoever, except its possible adventitious pre-
sence in the tray, and is a disregard of specimens that
Linnaeus himself documented as type. I ... see no reason
for denying to the specimens of Drupa cornus ROpING
their status as syntypes.”
This restriction of the type lot to the two specimens
“marked” by Linnaeus, if accepted, further serves to in-
validate Cernohorsky’s lectotype designation, and at the
same time would fix the Linnaean name upon the species
now known as Drupa cornus R6pinc, a form that is
generically distinct from the species that are today referred
to the genus Mancinella Link, to which genus the manci-
nella of LAMaRCcK and authors, not of LinnagEus, has long
been referred. It was this consideration that led Dr. Keen
to request the International Commission on Zoological
Nomenclature to declare that the M. mancinella of Lin-
NAEUS, 1758, be considered a species dubium and that
the type be designated as the M. mancinella of authors.
The oldest valid name for the latter appears to be “Vole-
ma’ alouina Répine (1798, p. 58; sp. 728) based upon
the figure in Martini & CHEMNITZ, Conchylien Cabinet,
vol. 3, Tafel 101, figs. 967, 968. The Roding name is older
than Purpura gemmulata of Lamarck, 1816, based upon
the same illustration, which was listed by Dr. Keen as
the valid name (see Bulletin of Zoological Nomenclature,
vol. 21, prt. 6, p. 422).
The third specimen in the Linnaean collection, which is
completely unmarked by Linnaeus, is that which has been
selected by Cernohorsky as the lectotype. There is abso-
lutely no evidence that it was ever seen by Linnaeus, and
in fact, it seems most probable that it was subsequently
added during the interval between Linnaeus’ death in
1778 and the time of the Linnean Society’s acquisition of
the collection in 1828.
Vol. 12; No. 3
THE VELIGER
Page 369
It is pertinent to cite the history of the collection as
given by DopceE (1959, p. 179): “At the death of Lin-
naeus in 1778 his natural history collections, manuscripts
and library descended to his son who was himself a com-
petent naturalist and who had accumulated a collection
of his own. Based upon what we know of his abilities and
his conscientious and useful annotations in his own copy
of the twelfth edition of the Systema it is probable that
the elder Linnaeus’s Collections came to no harm during
the time they were in his son’s custody. Certainly there
is no evidence that any transfers of specimens were made
between the two collections. At the son’s death in 1783,
his mother, as his executrix, sold the elder Linnaeus’s
Collections and literary material to James Edward Smith
of London, then a young medical student. Smith was
knighted in 1814. The purchased material arrived in
London in 1784, and remained in Smith’s possession
until his death in 1828, when it was acquired by The
Linnean Society of London of which Sir James Smith
was the first President at its foundation in 1788 and re-
mained in that office until his death, All of the Linnaean
material is still in the possession and custody of the Society.
“Smith was primarily a botanist and, as might be ex-
pected, was less interested in the zoological portions of the
Linnaean Collections. It is admitted that during the many
years he held the collections the cabinet of molluscs, at
least, was mishandled through the inadvertent replace-
ment of specimens in the wrong receptacles as well as by
the integration of Smith’s own specimens. Many of the
equivocal situations now encountered in connection with
the undocumented specimens are the result of these un-
fortunate accessions ... .”
With respect to the mishandling of the collection while
in the possession of Sir James Smith, HANLEy states (1855,
p. 2): “Had the cabinet ... passed direct from the hands
of Linnaeus to the Museum of the Linnean Society, ... ,
without any intermediate possession by a third party, the
sole additions to the typical examples coeval with the
date of his publications would have been those he had
acquired subsequently to the appearance of the last edi-
ion of his ‘Systema;’ and as these were almost invariably
found packed in separate papers or pill-boxes (often in-
deed indicated as undescribed), and not distributed like
the original types, in metallic receptacles, their presence
would have caused but little impediment to any deter-
mined search. The collection, however, did not reach the
Society direct, but was held for a while by the noted bot-
anist Sir James Smith, during whose custody numerous
other specimens were mingled with the ancient ones. This
ill-advised admixture has not merely augmented to an
- almost inconceivable degree the difficulties of investiga-
tion, but has too frequently been fatal to any accurate
decision.”” HaNLEy also adds (p. 3) “The original speci-
mens, when large enough to permit it, had been inscribed
by Linnaeus either with their names or with numerals cor-
responding to their position in his ‘Systema;’ the smaller
ones had been deposited in tin boxes, marked in like
manner; oftentimes, indeed the numerals were written
on both shell and boxes.”
Thus all authors (HANLEY, SmitH, DopcE) who have
worked carefully with the Linnaean collection agree that
the original Linnean specimens were marked by Linnaeus.
In the present case the only specimens thus recognized
were the two examples of Drupa cornus Répinc; the
third specimen, unmarked, was thus either added sub-
sequent to the publication of the twelfth edition or, more
probably, by Sir James Smith as a typical example of the
“Murex” mancinella of Lamarck and other authors a-
mong his contemporaries,
It may be significant that the number on the two
marked specimens in question is that of the position of
the species in the twelfth edition, although the species
itself was described, under the number 469, in the tenth
edition. According to Haney (p. 3), “It is worthy of
remark, that these numerals more frequently correspond
to the series of the tenth than of the twelfth edition, a
fair ground for believing that these shells were not merely
his erroneous after-impressions of his own species, but
were admitted and recognized types when the final edi-
tion was printed.” Dopcr (1952, p. 8) more specifically
states that: “Where the number is of the tenth edition it
indicates that the specimen was a recognized and accepted
type when the twelfth edition was published.”
The fact that the number on the specimens of Murex
mancinella is that of the twelfth edition is grounds for the
suggestion that Linnaeus did not himself possess a speci-
men of this species at the time of the original description
in the tenth. It is known that some species which Linnaeus
did not himself possess were described from specimens
borrowed from the collections of Spengler, Count Tessin,
De Geer, Gyllenborg, and others (Dopce, 1959, p. 173),
and it may well be that the true “type specimen” of M.
mancinella was from one of these other sources.
The original description of Murex mancinella in the
tenth edition (1758, p. 751) is as follows: “M. testa ecau-
data ovata spinis obsoletis, apertura edentula, columella
transversim striata.”
While CerNonHorsky correctly points out that the spe-
cimens of Drupa cornus do not agree with this description
in that the aperture is not edentulous but is marked by
“5 - 6 prominent labial denticles”, he fails to note that the
M. mancinella of LAMaRcK and subsequent authors also
does not agree with the description for the columella is
smooth, not transversely striated, and the phrase “spinis
Page 370
obsoletis” does not conform to the sharp and prominent
spines seen in that shell. The first of these differences was
noted by HANLey (1855, pp. 295 - 296) and commented
upon in some detail by SmMrru (1913, p. 287) in the paper
cited by CerNoHorsky. SMITH stated: “The Purpura
mancinella of authors does not agree with Linné’s descrip-
tion in the tenth edition of the Systema, for the columella
is not ‘transversim striata’, and nothing, moreover, is
said as regards colour. The ‘apertura edentula’ is fairly
descriptive, for the red thread-like lines within the mouth
could hardly be termed teeth, yet one would expect such
a conspicuous feature to have been referred to if Linné
had the shell before him at the time.” The latter italics
are those of SmitH, who clearly did not believe that the
third, unmarked specimen was a part of the original Lin-
nean collection. Smiru also noted, as did HANLEy, that
the Murex mancinella of the Museum Ulricae of Linnae-
us, published in 1764, is “certainly, in part, the mancinel-
la auctorum, for this is shown by the ‘spinae brevissimae
purpurascentes’, which is a characteristic feature of that
species. The ‘fauce lutea, transversim striata’ [throat saff-
ron-yellow, transversely striated] also seems to indicate
this species” (SmiTH, 1913, p. 287).
The fact that Linnaeus returned to the original tenth
edition description of Murex mancinella in the twelfth
edition of the Systema without any additions or altera-
tions based upon the specimens in the Museum Ulricae
which he had studied in the interim, coupled with his
acceptance of the specimens of Drupa cornus as repre-
senting his species seems to the writer to be rather com-
pelling evidence that his concept was of a broad and
variable form that probably would include a majority
of the presently known species of Thaidinae, and hence,
as originally suggested by Smrru, the species should be
considered as a species dubium, an action that has been
requested of the International Commission by Miss Keen.
To summarize: The writer is of the opinion that all
available evidence strongly suggests (1) that none of the
specimens in the Linnaean collection can properly be
taken as representing the types of the Murex mancinella
Linnaeus, 1758, the two “marked” specimens bearing
the number of the species in the twelfth edition probably
having been added to his collection subsequent to 1758,
as suggested by CerNoHorRSKy, and the third unmarked
specimen probably having been inserted by Sir James
Smith in the interval between 1784 and 1828. Hence
THE VELIGER
Vol. 12; No. 3
neither the DopcE (1957) designation of the two marked
specimens as “syntypes” nor the CeRNoHoRSKY (1969)
designation of the third, unmarked one as the “lectotype”
can be considered as valid actions under the Code.
(2) Neither the Murex mancinella of authors nor the
“marked” specimens of Drupa cornus fit the original de-
scription, which, on the other hand, is too inadequate to
permit its restriction to any of the known species of Thai-
dinae and the name Murex mancinella of LINNAEUS,
1758, should be considered a nomen dubium. (3) The
correct name for the M. mancinella of Lamarck, 1822,
and subsequent authors must be taken as Mancinella
alouina (RO6pinc, 1798).
LITERATURE CITED
CERNOHORSKY, WALTER OLIVER
1969. The Muricidae of Fiji — Part II. Subfamily Thaidinae.
The Veliger 11 (4): 293-315; plts. 47-49; 21 text figs.; 1
map (1 April 1969)
Dopce, HENRY
1952. _A historical review of the mollusks of Linnaeus. Part 1.
The classes Loricata and Pelecypoda. Bull. Americ. Mus.
Nat. Hist. 100: 1 - 263
1957. _ A historical review of the mollusks of Linnaeus. Part 5.
The genus Murex of the class Gastropoda. _ Bull. Amer. Mus.
Nat. Hist. 113 (2): 77 - 224 (30 September 1957)
1959. _ Evidential factors in the identification of the Linnaean
molluscs. Journ. Linn. Soc. London, Zoology 44 (296) :
170 - 179 (April 1959)
HANLEy, SYLVANUS
1855. Ipsa Linnaei Conchylia; the shells of Linnaeus, deter-
mined from his manuscripts and collections. London, 556
pp.; 6 plts.
LINNAEus, CAROLUS
1758. Systema naturae per regna tria naturae ... editio
decima, reformata 1 [Regnum animale]. Stockholm (Laurentii
Salvii) pp. 1 - 824+1- iii
1767. | Systema naturae per regna tria naturae ... editio
duodecima, reformata 1 [Regnum animale] (2): 533 - 1327.
Stockholm (Laurentii Salvii)
Ropinc, PETER FRIEDRICH
1798. Museum Boltenianum sive catalogus cimeliorum
pars secunda continens Conchylia Hamburg, viii+
199 pp. (10 September 1798)
SmirH, Epcar ALBERT
1913. Note on Murex mancinella Linn. Proc. Malacol.
Soc. London 10 (4): 287 - 289 (28 March 1913)
Vol. 12; No. 3 THE VELIGER Page 371
A List of Recommended Nomenclatural Changes
for MAcFARLAND’S
“Studies of Opisthobranchiate Mollusks
of the Pacific Coast of North America’’
BY
RICHARD A. ROLLER
1127 Seaward Street, San Luis Obispo, California 93401
MANY WORKERS IN THE FIELD of opisthobranchiate mol-
lusks have felt the need for a review of the nomenclature
of the species treated in MacFarLanp’s posthumously
published monograph. Because of restrictions relating to
the MacFarland estate, the editors were prevented from
making changes at the time of publication by the Califor-
nia Academy of Sciences, and the monograph was pub-
lished using MacFar.anp’s original names.
Some of this needed work has been done already in
various publications (see: STEINBERG, 1961, 1963; Opx-
cus, 1967; Burn, 1968; SpHon «& Lance, 1968; and
Rotter, 1969). The present list, prepared at the sugges-
tion of the Department of Invertebrate Zoology of the
California Academy of Sciences, brings together all of the
recommended nomenclatural changes, old and new.
The animals treated in the following list are in the order
of their appearance in the monograph. The names used
by MacFar.anp appear in the left column (those pro-
posed as new in the monograph without author), while
the recommended name changes are presented in the right
NER, 1963; Lance, 1966; Franz, 1967; Marcus « Mar-
MacFartanp Names
Aglaja diomedea (plate 2, figure 4)
Aclesia rickettsi
Il I
Hermaea ornata (BErcH)
Phyllobranchopsis enteromorphea [sic] CocKERELL &
Extot, 1905
Elysia bedeckta
Pleurobranchus californicus DALL
Acanthodoris columbina MacF ar anb, 1926
Corambella bolini
Glossodoris macfarlandi (CocKERELL)
Chromodoris californiensis BERGH
Chromodoris porterae COCKERELL
Austrodoris odhneri
Petelodoris spongicola
Dendrodoris fulua (MacFaruanp, 1905)
column.
New Names
Aglaja ocelligera (BERGH, 1894)
Stylocheilus longicauda (Quoy & Gatmarp, 1824) (see
BEEMAN, 1968)
Placida dentritica (ALDER & Hancock, 1843) (see Lone,
1969
Hermaeina smitht Marcus, 1961
Elysia hedgpethi Marcus, 1961 (s.SpHON & LANcE, 1968)
Berthella californica (Dati, 1900) (see Lance, 1966)
Acanthodoris nanaimoensis O’DonocHuE, 1921 (see
STEINBERG, 1963)
Doridella steinbergae (LANCE, 1962) (see FRANz, 1967)
Chromodoris macfarlandi CockERELL, 1902
Hypselodoris californiensis (BERGH, 1879)
Hypselodoris porterae (CocKERELL, 1902)
Archidoris odhneri (MacFar.anp, 1966) (s. Burn, 1968)
Atagema quadrimaculata CoLuiER, 1963
Doriopsilla albopunctata (Cooper, 1863) (see STEINBERG,
1961)
Page 372
Dendrodoris albopunctata (CoorER)
Duvaucelia tetraquetra (PALLAS)
Duvaucelia festiva (STEARNS)
ll Il
Duvaucelia exsulans (BERGH)
Duvaucelia gilberti
|
Dendronotus venustus
Doto varians (C. group of dark- colored specimens)
Antiopella aureocincta
Coryphella fisher
Eubranchus occidentalis
Cuthona rosea
Wl ll
Cratena rutila
Cratena flavovulta
Cratena fulgens
Cratena albocrusta
Cratena virens
Cratena abronia =
Cratena spadix
Phidiana nigra
Aeolidiella oliviae
In treating Aglaja, MacFarLanp described several lots
of specimens. Most of these appear to be assigned to the
correct species, A. diomedea (Brercu, 1894), according to
the shell figures shown (plt. 6, fig. 8; plt. 7, figs. 11-14).
However, a specimen received by him from M. W. Willi-
ams in 1950 (described on pp. 8-9) was incorrectly as-
signed to A. diomedea. The caudal lobes of that specimen
were described as having “The left lobe, triangular, of
about the same size as the right one but is prolonged far
beyond it as a delicate filament (flagellum) ... The
filament is 2.5 to 3mm in length, extending beyond the
posterior foot margin.” No description of the shell was
given. MacFar.anb states that “The specimen matched
the colored figure (pl. 2, fig. 4) of the original specimen
_ taken from Monterey Bay-in 1894.” The plate descrip-
tion on page 396 describes the animal as having the left
caudal lobe lengthened into a slender flagellum, and the
colored figure on plate 2 shows this flagellum clearly.
THE VELIGER
Vol. 12; No. 3
Doriopsilla albopunctata (Coorrr, 1863)
Tochuina tetraquetra (Pautas, 1788) (s. ODHNER, 1963)
Tritonia festiva (STEARNS, 1873) (s. Marcus & Marcus,
1967)
Tritonia exsulans BercH, 1894 (see Marcus & Marcus,
1967)
Tritonia gilberti (MacFarLanp, 1966) (see Marcus &
Marcus, 1967)
Dendronotus frondosus (AscANIuS, 1744)
Doto kya Marcus, 1961
Antiopella barbarensis (Cooper, 1863)
Lance, 1968)
Coryphella trilineata O’DoNocHUE, 1921
Capellinia rustya Marcus, 1961
Precuthona divae Marcus, 1961 (see SPHON & LANCE,
(see SPHON &
1968)
Trinchesia lagunae (O’DonocHuE, 1926) (see ROLLER,
1969)
Trinchesia flavovulta (MacFartanp, 1966) (see ROLLER,
1969)
Trinchesia fulgens (MacFartanp, 1966) (see ROLLER,
1969)
Trinchesia albocrusta (MacFar.anp, 1966) (see ROLLER,
1969)
Trinchesia virens (MacFartanp, 1966) (see ROLLER,
1969)
Trinchesia abronia (MacFar.anpD, 1966) (see ROLLER,
1969)
Catriona alpha (Basa & Hamartant, 1963) (see ROLLER,
1969)
Phidiana pugnax Lance, 1961 (s. SpHoN & Lance, 1968)
Spurilla oliviae (MacFartanp, 1966)( s. SPHON & LANCE,
1968)
BERGH (op. cit., pp. 211, 213) described Aglaja diome-
dea as having the left lobe without flagellum; and A.
ocelligera as having the left lobe extended intoa 1 mm long
flagellum. The text description of the Williams specimen
and the plate description and figure of the 1894 Monte-
rey specimen indicate a flagellum. Therefore these 2 spe-
cimens should have been identified as Aglaja ocelligera
(Bercu, 1894).
The naming of a new subspecies, Pleurobranchus cali-
fornicus denticulatus by MacFaruanp presents a difficult
nomenclatural problem. Lance (1966) transferred Pleu-
robranchus californicus Dati, 1900 to Berthella califor-
nica (Dax, 1900) on the basis of the modern separation
of the genera of the family Pleurobranchidae according
to the configuration of the gill rhachis. In Berthella BLaIn-
VILLE, 1825, the gill rhachis is smooth; while in Pleuro-
branchus Cuvier, 1805, the gill rhachis bears a series of
tubercles at the junction of the gill plumules with the
Vol. 12; No. 3
THE VELIGER
Page 373
rhachis. If Datt’s original description of the species “The
gill short, its stem finely granular, not tuberculate ...”
is accurate, then BercH’s (1902) anatomical account
of the same specimen, wherein he states “... the plume
made up of alternating tubules at the base of the pinnules,
owing to the dilation of the branchial veinules as they join
the branchial vein ...,” is somewhat contradictory. If,
in fact, DALL’s species has a smooth gill rhachis, then it
most properly belongs in Berthella, as suggested by LANCE
(op. cit.). However, since the gill rhachis of P californicus
denticulatus is described as tuberculate, MAcFARLAND’s
species must be placed in Pleurobranchus, and cannot re-
main a subspecies.
The proper status of Pleurobranchus californicus denti-
culatus depends upon a better understanding of the ana-
tomy of P californicus s.s. Further study of live material
from the type localities of both taxa will be required be-
fore this problem can be satisfactorily solved.
MacFar.anp also named another pleurobranchid in
his monograph, Pleurobranchus strongi. This species was
described as having the “ rachis smooth, without
tubercles ... ,” and on this basis should be placed in
Berthella. However, he also observed a pedal gland and
a prostate gland in his material of P strongi; and these
characteristics should not be present in members of the
genus Berthella, according to BurN’s synoptic key to the
Pleurobranchacea (1962). Macnae (1962) does not con-
sider these characteristics to be of value for generic sep-
aration. The proper generic placement of this species is
dependent upon further study of live material from the
type locality and a much needed revision of the family
Pleurobranchidae.
Another nomenclatural problem exists due to the
naming of Doto varians. MAcFaRLAND stated that of the
many specimens collected by him over a period of years,
“An effort has been made to isolate these into groups
based largely upon coloration. Three distinct groups are
recorded.” He listed the three groups as: A. The speci-
mens of lightest colorings; B. The yellow specimens;
and C. Group of dark-colored specimens, and stated that
“Intermediate forms between the light and dark varieties
of coloration are rare, and a complete graduation has
never been found.”
OpHNER (1936), in his discussion of the nomenclature
of the genus Doto OxEN, 1815, stated that “The classifi-
cation of the about 20 species which prove valid, offers
great difficulties, since only slight differences exist and
these are above all to be found in the colouration, ... A
classification of Doto must base upon this character,
at least for the present.” He rejected as useful character-
istics for species separation the use of radular teeth and
gills on the inside of the cerata.
Marcus (1961) named 4 species of Doto from the
Monterey Bay area, namely D. amyra, D. ganda, D. kya,
and D. wara. Of these species, only D. kya has any dark
pigment mentioned, while the 3 remaining species are all
described as light-colored.
The group of Doto specimens which MacFar.Lanp de-
scribed as “dark-colored” and figured on plate 42 (figures
2, 4, 8) are similar in coloration to D. kya, and should be
considered as belonging to that species. However, the
“lightest coloring” and “yellow” specimens of MAcFar-
LAND are not well enough described to allow allocating
them to any of the 3 light-colored species of Marcus
(1961). There is much work needed to be done on the
members of this: genus before this species (D. varians)
can be properly named.
There is a typographical error on page 51 in the de-
scription of Elysia bedeckta. In the last paragraph on that
page, the second sentence states that “The cusp has a
length from its anterior basal projection to the tip, of
0.33 mm ...” This figure should be 0.033 mm, according
to the original descriptive notes of MacFarland.
I wish to thank Mr. Allyn G. Smith, of the California
Academy of Sciences, for suggestions and critical com-
ments made during the preparation of the manuscript.
LITERATURE CITED
BEEMAN, RosBErRT Davip
1968. Order Anaspidea.
prt. 2) :87- 102; plt. 11; 12 text figs.
Bercy, Lupwic SopHus RupoLtpH
1894. Reports on the dredging operations off the west coast of
central America to the Galapagos, to the west coast of
Mexico and in the Gulf of California, in charge of Alexander
Agassiz, carried on by the U. S. Fish Commission steamer “Alba-
tross’, during 1891. XIII. Die Opisthobranchien. _—_ Bull. Mus.
Comp. Zool., Harvard Univ. 25 (10): 125 - 233; plts. 1-12
(October 1894)
1902. Malakologische Untersuchungen in Semper’s Rei-
sen im Archipel der Philippinen. Vol. 7, Abt. 4, Abs. 4:
313 - 382; plts. 25 - 29.
The Veliger 3 (Supplement,
(1 May 1968)
{not seen}
1904. | Nudibranchiata, Tectibranchiata-Pectibranchiata. In:
Semper, Reisen im Archipel der Philippinen. 9 (6), Lief. 1:
1-55; plts. 1-4 {not seen}
Burn, RoBERT
1962. On the new pleurobranch subfamily Berthellinae (Mol-
lusca: Gastropoda) ; a revision and new classification of the
species of New South Wales and Victoria. Mem. Nat. Mus.
Melbourne, no. 25: 129-148; plts. 1, 2; figs. 1-5
(1 May 1962)
1968. Archidoris odhneri (MacFartanp, 1966) comb. nov.,
with some comments on the species of the genus on the
Pacific Coast of North America. The Veliger 11 (2):
90 - 92 (1 October 1968)
Page 374
THE VELIGER
Dati, Witt1amM HEALey
1900. A new species of Pleurobranchus from California.
Nautilus 14 (11): 125-126
Franz, Davp R.
1967. On the taxonomy and biology of the dorid nudibranch
Doridella obscura. The Nautilus 80 (3): 73-79; 1 fig.
Lance, JAMES ROBERT
1966. New distributional records of some northeastern Pacific
Opisthobranchiata (Mollusca: Gastropoda) with descriptions
of two new species. The Veliger 9(1): 69-81; 12 figs.
(1 July 1966)
Lone, STEVEN J.
1969. Records of Trinchesia virens, Trinchesia fulgens, and
Placida dentritica from San Luis Obispo County, California.
The Tabulata 2 (4): 9-12; 2 figs. (1 October 1969)
MacFaruanp, Frank Mace
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
Macnag, WILLIAM
1962. Notaspidean opisthobranchiate molluscs from South Afri-
cas Ann. Natal Mus. 15 (15): 167-181; 7 figs.
Marcus, ERNST
1961. | Opisthobranch mollusks from California. The Veli-
ger 3 (Supplement, pt. 1): 1-85; plts. 1 - 10.
(1 February 1961)
Marcus, EvELINE & ERNsT Marcus
1967. American opisthobranch mollusks. Studies in tropical
Oceanography no. 6. Inst. Marine Sci., Univ. Miami,
Florida; viiit+ 256 pp.; 1 plt.; 155+495 text figs. | (December)
Opune_rR, Nits HyjALMar
1936. | Nudibranchia Dendronotacea. A revision of the System.
Mélanges Paul Pelseneer. Mém. Mus. Roy. d’Hist. Nat.
de Belgique, Ser. II, Fasc. 3: 1057-1128; 1 plt.; text figs. 1-47.
1963. On the taxonomy of the family Tritoniidae (Mollusca:
Opisthobranchia). The Veliger 6 (1): 48-52 (1 July ’63)
Ro ier, RicHarp A.
1969. | Nomenclatural changes for the new species assigned to
Cratena by MacFartanp (1966). The Veliger 11 (4):
421-423; 2 figs. (1 April 1969)
Rotier, RicHArD A. & STEVEN J. Lone
1969. An annotated list of opisthobranchs from San Luis Obis-
po County, California. The Veliger 11 (4) : 424-430; 1
map (1 April 1969)
SPHON, Gate G., Jr., x JAMES RoBerT LANCE
1968. An annotated list of nudibranchs and their allies from
Santa Barbara County, California. Proc. Calif. Acad. Sci.
(4) 36 (3): 73-84; 1 fig. (25 September 1968)
STEINBERG, JOAN EMILY
1961. Notes on the opisthobranchs of the west coast of North
America. — I. Nomenclatural changes in the order Nudi-
branchia (Southern California). | The Veliger 4 (2) : 57 - 63.
(1 October 1961)
1963. Notes on the opisthobranchs of the west coast of North
America. III. Further nomenclatural changes in the order
Nudibranchia. The Veliger 6 (2): 63-67 (1 Oct. 1963)
Vol. 12; No. 3
Vol. 12; No. 3
NOTES & NEWS
Range Extensions for
Acanthodoris hudson’ MACFARLAND, 1905,
and Onchidoris bilamellata (LINNAEUS, 1767)
BY
GARY R. McDONALD
310 Graves Street, San Luis Obispo, California 93401
On May 31, 1969, a single specimen of Acanthodoris
hudsoni MacFarxanp, 1905 which measured 11 mm in
length and 84 mm in width was collected by Mr. Hans
Bertsch from the boat docks at Morro Bay, California.
On June 3, 1969, a second specimen, measuring 24 mm in
length and 9 mm in width was collected by the author in
the low tide zone just south of Spooner’s Cove, Montana
de Oro State Park, San Luis Obispo County, California.
On July 11, 1969, a third specimen (16 mm long, 11 mm
wide) was collected from a depth of 20 feet at Shell
Beach, also in San Luis Obispo County. To the author’s
knowledge these are the first recorded collections of this
species south of Monterey Bay, California, a range ex-
tension of about 125 miles.
On May 16, 1969, a single specimen of Onchidoris bi-
lamellata (Linnagus, 1767), measuring 9 mm in length,
was collected by the author from the boat docks at Morro
Bay, California. On June 26, 1969, 4 more specimens
which measured 20, 20, 22, and 25 mm in length were
collected from buoy no. 7 in Morro Bay. On July 24, 1969,
2 more specimens (11 mm and 16 mm long) were collec-
ted from a depth of 20 feet in Morro Bay. To the author’s
knowledge these are the first recorded collections of this
species south of Monterey Bay, California, a range ex-
tension of 110 miles.
All identifications were made or verified or both by
Mr. Richard A. Roller of San Luis Obispo, California.
REFERENCES CONSULTED
Lance, JAMES ROBERT
1961. A distributional list of Southern California Opistho-
branchs. The Veliger 4 (2): 64-69 (1 October 1961)
THE VELIGER
Page 375
MacFar.anb, Frank Mace
1905. A preliminary account of the Dorididae of Monterey
Bay, California. Proc. Biol. Soc. Washington, 18: 35 - 54
Marcus, Ernst
1961. | Opisthobranch mollusks from California. The
Veliger 3 (Supplement, pt. I): 1-85; plts. 1-10. (Feb. 1, 1961)
Rotter, RicHarp A. & STEVEN J. Lone
1969. An annotated list of opisthobranchs from San Luis Obis-
po County, California. The Veliger 11 (4): 424-430; 1
map (1 April 1969)
SpHon, Gate G., Jr., « JamES RoBERT LANCE
1968. An annotated list of nudibranchs and their allies from
Santa Barbara County, California. Proc. Calif: Acad. Sci.
(4) 36 (3): 73-84; 1 fig. (25 September 1968)
STEINBERG, JoAN EmILy
1963. Notes on the opisthobranchs of the West Coast of North
America - IV. A distributional list of opisthobranchs from Point
Conception to Vancouver Island. The Veliger 6 (2): 68 to
73 (1 October 1963)
Spawning Notes V.
Acanthina angelica 1. OLDROYD, 1918
and Acanthina lugubris (SOWERBY, 1821)
BY
FAY HENRY WOLFSON
Associate Curator of Marine Invertebrates
San Diego Natural History Museum, San Diego
California 92112
(6 Text figures)
FROM MID-WINTER THROUGH late spring, the egg capsules
of Acanthina angelica 1. OtpRoyp, 1918 are frequently
found in the high intertidal zone throughout the northem
Gulf of California. I have observed the spawning process
at Bahia de los Angeles from January through April in
1968 and 1969, at Puerto Refugio in Isla Angel de la
Guarda in December 1967, and on Isla Cardonosa in
April 1969. Fresh capsules were seen by Carol Skoglund
(personal communication) at Cholla Bay in March and
April 1969 and by me at Bahia San Luis Gonzaga in May
1968. In January 1969, I collected a cluster of capsules
and two spawning females at Playa Alicia, south of San
Felipe.
Page 376 THE VELIGER Vol. 12; No. 3
Table 1
Acanthina angelica Acanthina lugubris
Capsule
Average maximum dimensions 6.66 X 2.49 mm 8.64 X 3.94 mm
Color (relative) Pale yellow Bright yellow
Relative thickness of walls Thin Thick
Stalk Cylindrical Flattened cylinder
Eggs
Average number per capsule more than 200 less than 200
Average diameter 119 p 157 p
Position of egg mass! Above the stalk at base of capsule, occu- Loosely filling entire length of capsule, but
pying up to 4 of the interior space not touching the inner capsule walls
Egg mass! Enclosed in a thin, transparent membrane, The eggs form 2 or more irregular columns,
attached in one or more areas to capsule each column several eggs deep at the cen-
walls ter. The underlying eggs are embedded in
a spongy matrix, which also forms a flat
sheet separating the columns. This sheet
appears to consist of a single layer of cells
which resemble flattened eggs. The whole
mass lies free within the capsule
1 see schematic section
Figure 3
Acanthina lugubris (SowERBy, 1821)
Schematic section through an egg capsule
Figure 1
Acanthina lugubris (SowERBy, 1821)
Egg cluster. Line indicates where section was made for Figure 3
5mm
Figure 2 Figure 4
Acanthina lugubris (SowErsy, 1821) Acanthina angelica 1. OtpRoyp, 1918
A single egg capsule seen from the top Egg cluster. Line indicates where section was made for Figure 6
The capsules present an interesting comparison with 1821). One of these is with SDNHM lot no. 22722, col-
two clusters of the spawn of Acanthina lugubris (SowErRBy, lected by Henry Hemphill on Isla Cedros and presented to
Vol. 12; No. 3
the museum by Herbert Lowe. The other is a cluster
which I collected on 12 March 1968 on the westernmost
island of the San Benito trio.
Figure 5
Acanthina angelica I. OtpRoyp, 1918
A single egg capsule seen from the top
Figure 6
Acanthina angelica I. OtpRoyp, 1918
Schematic section through an egg capsule
The Acanthina lugubris capsules I collected were being
deposited by 2 females in a crevice of a rocky cliff, high
above the low-tide line. In contrast, A. angelica typically
spawns on the undersides of rocks in groups of a dozen or
more individuals. Both the Gulf and coastal capsules are
of the characteristic muricid vase type, cemented to the
substrate by a fused basal membrane. Other compara-
tive data are presented in Table 1.
The drawings are by Anne Acevedo of the SDNHM.
ERRATA
The figures on Plate 27 of our October issue were, through
an error, numbered wrongly. Instead of Figures 8 to 13,
the numbers should be Figures 1 to 6. We apologize.
The Editor.
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THE VELIGER
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THE VELIGER
Page 379
METHODS & TECHNIQUES
A Mechanical Shell Washer
BY
JACQUELIN N. MILLER
Department of Oceanography, University of Hawaii
AND
CARL L. HUBBS
Scripps Institution of Oceanography
University of California, San Diego, California
MOLLUSCAN MATERIAL Collected from archeological sites
typically requires major cleaning before it can be identi-
fied to species or analyzed by weight. Frequently the
extraneous coating is particularly tenacious, apparently be-
cause the charcoal and soil fuse to the shell during
heating in midden fires and/or the encrusting caliche
becomes firmly cemented to the shells.
During the course of studies on the molluscan fauna
from ancient archeological sites (MILLER « Husss, MS
in preparation) need arose to devise a method more rapid
and thorough than the usual hand scrubbing technique
for cleaning the rapidly accumulating, voluminous quan-
tities of heavily soiled shells. To meet this need, the me-
chanical shell washer described below was developed (see
Figure 1).
Basically, the device is similar to the standard labora-
tory rock tumbler or jar mill, with certain modifications
to better adapt it to our purposes. The shell containers are
polyethylene cylinders 23 & 15cm with removable in-
ternal baffles constructed from 4-inch stock sheets of
lucite. The snap-on polyethylene lids are kept secure and
watertight by turned aluminum caps connected by screw
‘ Contribution from Scripps Institution of Oceanography, Uni-
versity of California, San Diego, California 92037
Page 380 THE VELIGER Vol. 12; No. 3
Figure 1
rods and wing nuts. ‘These cylinders, tightly sealed by the GALL,
caps, ride freely on two sets of rollers driven by a 1/15 hp. |<—— 15 cm ——>|
electric motor. A simple frame of stainless steel rods and
laboratory clamps attached to a heavy wooden base keeps
the cylinders centered on the rollers (Figure 2).
To wash a sample, the cylinders are filled approximately
3 full of shell. Hot water and 6 cc of a non-foaming de- (: if Loreen ff i
tergent are added to within 3 cm of the top. For shells =: |
heavily encrusted with caliche a few drops of hydrochloric
acid may be added. The polyethylene lids are then sealed
with the aluminum caps and the cylinders are placed on
the rollers to wash for 15 to 45 minutes, depending on the
size and condition of the sample. When a sufficient time
has elapsed, the cylinders are removed and the sample is
rinsed into a 4mm sieve. The soil passing through the
sieve is collected in a tray where it is available for further
examination. The cleaned shells remaining in the sieve
are oven-dried preparatory to sorting for species and
analysis by weight.
We feel that this particular machine has several ad-
vantages over the commercially available models. Use of
polyethylene in place of metal or glass cylinders assures
virtually no breakage of even the more fragile species.
Fewer problems of leakage are encountered with the
free-riding cylinders than with direct-drive cylinders. The
polyethylene cylinders are much lighter and more easily
handled in loading and unloading than comparable
sized metal, porcelain, or glass cylinders, and there is no
corrosion problem. The entire assembly, complete with
cylinders and labor, may be reproduced for about two-
thirds the cost of commercially available tumblers.
6 Polyethylene Cylinder
R ble Lucite Baffle Ey) yy
ertetee c 8 with Baffle inserted
i LLL
Aluminum Caps Complete Assembly
Figure 2
Vol. 12; No. 3
THE VELIGER
Page 381
BOOKS, PERIODICALS, PAMPHLETS
Superfamilia UNIONACEA
by Dr. Fritz Haas, Chicago, Illinois. x-+663 pp., 5 text
figs. Part 88 of “Das Tierreich,” publ. Walter de Gruyter
& Co. Berlin (in German). 380.- DM (= approximately
US$ 105.- at present exchange rates).
Dr. Haas has produced an exceedingly thorough sys-
tematic study of the pearly fresh-water mussels of the
world, which no doubt will serve as a basic reference for
many years to come. He is eminently qualified to under-
take such an extensive work, having already published on
the Bivalvia in Bronn’s well-known “Klassen und Ord-
nungen des Tierreichs” covering more than 1900 pages of
text. The statistically minded will be interested to learn
that the Unionacea is divided into 4 families, broken down
into 6 subfamilies, 142 genera, 111 subgenera, and 826
species (the considerable number of subspecies not de-
termined for this review).
Dr. Haas’ treatment includes several features that the
systematist will find very useful. Dichotomous keys to the
various subfamily, genus, subgenus, and species groups are
provided; type localities as well as geographic ranges are
given for each species in addition to brief diagnoses. Syn-
onymies are arranged chronologically along with complete
original references, making an over-all terminal biblio-
graphy unnecessary. Names in synonymy include the orig-
inal references only. No one can accuse the author of not
being a thorough bibliographer as indicated, for example,
by the 526 entries for Anodonta cygnea Linnaeus for the
period 1758 to 1945, occupying 21 text pages! There is
a good index.
According to Haas only 6 species of the Unionacea
occur in western North America. These are Margaritifera
margaritifera (LINNAEUS, 1758), Gonidea angulata (LEa,
1838), Anodonta (Anodonta) beringiana MippENDORFF,
1851, A. (A.) oregonensis Lea, 1838, A. (Arnoldina) dejec-
ta Lewis, 1875, and A. (Brachyanodon) impura Say,
1829. If this arrangement is adopted some name changes
will be necessary for certain well-known Pacific Coast
species. Anodonta wahlametensis Lea, 1838, A. nuttalliana
Lea, 1838, and A. nuttalliana idahoensis HEMPHILL, 1891,
are listed as synonyms of A. oregonensis; similarly, A.
californiensis Lea, 1852 becomes a synonym of A. impura.
The extent to which these taxonomic changes should be
accepted is left to specialists in the group.
__ The price of the volume, quoted at 380 Mark, is the
equivalent of over $100.- at the re-valued exchange rate.
This, unfortunately, is far beyond the means of many
scientists and scientific libraries and is extremely high for
a splendid scientific work that should be readily available
to all systematic biologists, especially the malacologists.
At current rates of 3 cents for Xerox copies per page, the
entire book could be reproduced for $20.19. Thus the
unusually high initial cost will no doubt lead to a clandes-
tine and frequent violation of the copyright restriction
placed on this book.
AGS
Pleistocene Molluscs from the Namaqualand Coast
by A. J. Carrincton &« B. F Kens ey. Ann. South
African Mus., Vol. 52, part 9, pp. 189 - 223; plts. 18 - 29;
11 text figs. July 1969.
Twenty species of mollusks of Pleistocene age are re-
ported and discussed from raised beaches on the Nama-
qualand coast of South Africa. Of these, 17 species and
subspecies are described for the first time. A new genus,
Namamurex, type by M, N. odontostoma, spec. nov. (p.
198; plts. 20, 21), is compared with Acanthina, Cerato-
stoma, Pterorytis and Jaton. The shell of this new taxon
has 4 or 5 whorls, 3 varices per whorl but no sculpture
between the varices. The aperture is longer than the spire.
There are about 8 nodules on the inner margin of the outer
lip and just below these a prominent tooth. The anterior
canal is open.
A species of interest to west American workers is Hes-
pererato oppenheimeri, spec. nov. (p. 199; plt. 19, fig. C),
which is compared with H. vitellina (Hinps, 1844), orig-
inally described from Magdalena Bay, Lower California,
Mexico.
The authors state that the generic affinities of the new
species and the geographic ranges of previously recorded
species in this assemblage suggest that during early Pleis-
tocene time, at least, the water along the adjacent coast
was considerably warmer than today.
LGH
Catalogo de los Moluscos Marinhos del Uruguay
by ALFREDO FicuEmRAS & Omar E. Sicarpi. = Communic.
Soc. Malacolég. Uruguay, Vol. 2, no. 15, pp. 255 - 275,
laminas 1 and 2 (pp. 274 - 275). October 1968.
This (the second) part of a catalog of the marine mol-
lusks of Uruguay includes 12 species of Scaphopoda and
36 species of Pelecypoda. Geographic distribution only is
reported for the Scaphopoda; descriptions, synonymy,
Page 382
THE VELIGER
Vol. 12; No. 3
comments and geographic distribution are given for the
bivalves, and all except 3 are illustrated by line drawings.
In addition, a line drawing of a single valve of each of
3 species of Amphineura is included.
LGH
Systematics, Ecology, and Distribution of the Mollusks
of Los Roques, Venezuela
Bull. Mar. Sci. (Univ. Miami
Sept. 1969.
by Ropert C. Work.
Press), Vol. 19, no. 3, pp. 614 - 711; 4 figs.
One hundred and twenty five species of marine mol-
lusks from San Roque, Venezuela, are distributed in 4
Classes: Amphineura (3), Gastropoda (87), Pelecypoda
(34), Cephalopoda (1). An historical summary is given,
and the zoogeography and ecology are discussed as well
as the feeding habits of some of the gastropods. Records
of the occurrences of the species are reported and a special
effort is made to establish accurate geographic ranges.
LGH
Contributions to the Knowledge
of South African Mollusca. Part VI. Supplement.
by K. H. Barnarp.
prt. 4, pp. 595 - 661, plts. 1, 2, 30 text figs.
Ann. South Afr. Museum, Vol. 47,
July, 1969.
This posthumous paper by KeppELL Harcourt Bar-
NARD was prepared for publication by B. EF Kens.ey.
About 156 species and 2 varietal forms are reported and
discussed, including notes on many other species. All are
gastropods, except 2 pelecypods. Of the total number, 10
forms are identified only as to genus. The radulae of many
of the gastropods are described or illustrated. Illustrations
include 22 species previously described or mentioned by
BaRNARD in earlier parts of a series of papers dealing with
the marine mollusks of South Africa. Families especially
well represented in the present paper are the Turritidae
(34 species, 1 variety, 5 others identified only as to genus)
and the Marginellidae (14 species).
The following 16 species are described for the first
time: Surcula aditus, Drillia dovyalis, D. pecus, D. erepta,
Pleurotoma curricula, Persicula nigrocrocea, P. alborubida,
Nassa rhysonepia, Trophon beatum, Tritonalia juritzi, T.
aedicularum, Fossarus translucens, Scala munda, ?Retro-
tortina cuniculus, Turbo boswellae, Calliostoma circus.
An extensive bibliography is included.
LGH
A Monograph of the Cephalopoda of the North Atlantic:
The Family Joubiniteuthidae
by Ricrarp E. Younc & CiypE F E. Roper. Smithson.
Contrib. Zoolog. no. 15: pp. 1 - 10; 6 text figs. U.S.
Govt. Printing Off., Washington, D.C. 20402, 25 cents.
1969.
In this beautifully illustrated paper the authors syn-
onymize Valdemaria danae Jounin, 1931 with Joubini-
teuthis portieri (Jounin, 1912) ; they confirm the validity
of the family Joubiniteuthidae as established by NaEF in
922%
RS
An Illustrated Key to the Families
of the Order Teuthoidea (Cephalopoda)
by Ciype E E. Roper, RicHarp E. Youne & Giipert L.
Voss. Smithson. Contrib. Zoolog. no. 13: pp. 1 - 32;
16 text figs. U. S. Govt. Printing Off., Washington, D.
C, 20402, 45 cents. 1969.
A dichotomous key leads to the 25 families of the order
Teuthoidea; illustrations include representatives of each
family; and the current state of systematics within each
family is discussed briefly.
RS
The Tertiary Non-Marine Mollusca of South America
by Juan J. Paropiz. Ann. Carnegie Mus., vol. 40 etc.
This excellent work was discussed in our October issue
(vol. 12, p. 236) ; we indicated at that time that we were
not able to inform our readers how this work might be
acquired. We are delighted to be able to rectify this now:
this scholarly work can be obtained by writing to Carnegie
Museum, 4400 Forbes Avenue, Pittsburgh, PA 15213
The book is priced at $7.- plus a 25 cent shipping charge.
Editor.
THE VELIGER is open to original papers pertaining to any problem concerned
with mollusks.
This is meant to make facilities available for publication of original articles
from a wide field of endeavor. Papers dealing with anatomical, cytological, distri-
butional, ecological, histological, morphological, physiological, taxonomic, etc.,
aspects of marine, freshwater or terrestrial mollusks from any region, will be
considered. Even topics only indirectly concerned with mollusks may be acceptable.
It is the editorial policy to preserve the individualistic writing style of the
author; therefore any editorial changes in a manuscript will be submitted to the
author for his approval, before going to press.
Short articles containing descriptions of new species or other taxa will be given
preferential treatment in the speed of publication provided that arrangements
have been made by the author for depositing the holotype with a recognized
public Museum. Museum numbers of the type specimens must be included in the
manuscript. Type localities must be defined as accurately as possible, with geo-
graphical longitudes and latitudes added.
Short original papers, not exceeding 500 words, may be published in the column
“NOTES and NEWS’; in this column will also appear notices of meetings of
regional, national and international malacological organizations, such as A.M. U.,
U. M.E., W.S.M., etc., as well as news items which are deemed of interest to
our Members and subscribers in general. Articles on “METHODS and TECH-
NIQUES” will be considered for publication in another column, provided that
the information is complete and techniques and methods are capable of duplication
by anyone carefully following the description given. Such articles should be mainly
original and deal with collecting, preparing, maintaining, studying, photographing,
etc., of mollusks or other invertebrates. A third column, entitled “INFORMA-
TION DESK,” will contain articles dealing with any problem pertaining to
collecting, identifying, etc., in short, problems encountered by our readers. In
contrast to other contributions, articles in this column do not necessarily contain
new and original materials. Questions to the editor, which can be answered in this
column, are invited. The column “BOOKS, PERIODICALS, and PAMPHLETS”
will attempt to bring reviews of new publications to the attention of our readers.
Also, new timely articles may be listed by title only, if this is deemed expedient.
Manuscripts should be typed in final form on a high grade white paper, not
exceeding 81,” by 11”, at least double spaced and accompanied by a clear carbon
or photo copy. A pamphlet with detailed suggestions for preparing manuscripts
intended for publication in THE VELIGER is available to authors upon request.
A self-addressed envelope, sufficiently large to accommodate the pamphlet (which
measures 51,” by 81”’), with double first class postage, should be sent with the
request to the Editor.
EDITORIAL BOARD
Dr. Donatp P. Asgort, Professor of Biology
Hopkins Marine Station of Stanford University
Dr. Jerry DonouuE, Professor of Chemistry
University of Pennsylvania, Philadelphia, and
Research Associate in the Allan Hancock Foundation
University of Southern California, Los Angeles
Dr. J. Wyatr Duruam, Professor of Paleontclogy
University of California, Berkeley
Dr. E. W. Facer, Professor of Biology
Scripps Institution of Oceanography, La Jolla
University of California at San Diego
Dr. Caner Hanp, Professor of Zoology and
Director, Bodega Marine Laboratory
University of California, Berkeley
Dr. G Datias Hanna, Curator
Department of Geology
California Academy of Sciences, San Francisco
Dr. Jorn W. Hepcretu, Resident Director
Marine Science Laboratory, Oregon State University
Newport, Oregon
Dr. Leo G. HERTLEIN,
Curator of Invertebrate Paleontology
California Academy of Sciences, San Francisco
EDITOR-IN-CHIEF
Dr. RupoLF STOHLER, Research Zoologist
University of California, Berkeley
Dr. A. Myra KEEN, Professor of Paleontology and
Curator of Malacology
Stanford University, Stanford, California
Dr. Victor LoosanorF, Professor of Marine Biology
Pacific Marine Station of the University of the Pacific
Dr. Joun McGowan, Associate Professor of
Oceanography
Scripps Institution of Oceanography, La Jolla
University of California at San Diego
Dr. Franx A. Pirevxa, Professor of Zoology
University of California, Berkeley
Mr. Attyn G. Situ, Associate Curator
Department of Invertebrate Zoology
California Academy of Sciences, San Francisco
Dr. Ratpu I. Smiru, Professor of Zoology
University of California, Berkeley
Dr. Cuar.es R. Stasex, Associate Professor
of Zoology
Florida State University, Tallahassee, Florida
Dr. Donatp M. Witson, Professor of Biology
Department of Biological Sciences
Stanford University, Stanford, California
ASSOCIATE EDITOR
Mrs. JEAN M. Cate
Los Angeles, California
A Quarterly published by
CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC.
Berkeley, California
VOLUME 12 April 1, 1970
ConTENTS
A Report on the Feeding of Dendronotus iris on the Anthozoan Cerianthus sp. from
Monterey Bay, California (Plates 55 to 57)
Don R. Wosser . Rarity eh vcvel ob sate anne
Cargoa cupella, New Genus and New Species of Nudibranch from Chesapeake Bay
and the Generic Status of Okenia MENxKeE, Idalia LEucKarT, and Idalla
ORSTED (5 Text figures)
RosauizE M. VocEL & Leonarp P ScHULTz .
Malacological Applications of Scanning Electron Microscopy. I. Introduction and
Shell Surface Features (Plates 58 to 60)
ALAN SOLEM ay rey ry Ween aera en
The Effect of Wave Impact on some Aspects of the Biology of Sea Mussels
(9 Text figures)
J. R. E. Harcer .
Supplementary Comments on Deep Water Volutidae from the South China Sea and
South Africa (Plate 61)
Haratp A. REHDER . owe
Uptake of Sea Water into the Fluid Spaces of the Prosobranch Gastropod, Acmaea
scutum
H. H. WEBBER
Notes on the Deep Water Calliostomas of the Panamic Province, with Descriptions
of Six New Species (Plate 62)
James H. McLean . RNS os ls
An Annotated Bibliography of References to Marine Mollusca from the Northern
State of Sonora, Mexico
Caro, SKOGLUND
[Continued on Inside Front Cover]
- 383
. 388
- 394
- 415
- 417
. 421
EEE
Distributed free to Members of the California Malacozoological Society, Inc.
Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc.
Volume 13: $18.- Domestic; $19.- in the Americas; $19.50 in all other Foreign Countries
Single copies this issue $14.-. Postage extra.
Send subscription orders to Mrs. JEAN M. Care, 12719 San Vicente Boulevard,
Los Angeles, California 90049. Address all other correspondence to Dr. R. STOHLER, Editor,
Department of Zoology, University of California, Berkeley, California 94720
Second Class Postage paid at Berkeley, California
Contents — Continued
The Systematics and Some Aspects of the Ecology of the Genus Dendronotus
(Gastropoda : Nudibranchia) (Plates 63 and 64; 28 Text figures)
Gorpvon A. RoBILLIARD . Na ee PE CU NLn ee ics cy 0G. |e) aod!
- 433
Notes on the Egg Capsules and Larval Development of Conus purpurascens BRODERIP
(10 Text figures)
James NYBAKKEN 10160225000. Bop.) Ie al ene e ee to oO
A Supplement to the Annotated List of Opisthobranchs from San Luis Obispo
County, California
RIGHARD’ A.) ROLLER of. 7. ss, ese cout on non at om OS
NOTES & NEWS... 88) (5 20) cA eee ree 484
Concerning the Validity of the New Species of Paravitrea Proposed by
BRANSON & BATCH, 1970. Lanpon T. Ross & LAwRENCE G. ABELE
BOOKS, PERIODICALS & PAMPHLETS............. . 487
Xs)
we,
eee
Note: The various taxa above species are indicated by the use of different type styles
as shown by the following examples, and by increasing indentation.
ORDER, Suborder, DIVISION, Subdivision, SECTION,
SUPERFAMILY, FamiLy, Subfamily, Genus, (Subgenus)
New Taxa
Vol. 12; No. 4
THE VELIGER
Page 383
A Report on the Feeding of Dendronotus iris
on the Anthozoan Cerianthus sp. from Monterey Bay, California
BY
DON R. WOBBER
San Francisco State College, San Francisco, California
(Plates 55 to 57)
INTRODUCTION
Dendronotus iris Cooprr, 1863, 1s FOUND subtidally
throughout the year in Monterey Bay, California, feeding
on the anthozoan Cerianthus sp. The association which
has evolved between the two animals has not been pre-
viously described and might be viewed as a simple preda-
tor-prey relationship, were it not for the fact that D. iris
crops but does not, or perhaps can not, destroy the
Cerianthus. After consuming only a very limited part of
this prey, the nudibranch moves to other Cerianthus in-
dividuals for further feeding. It is rare in nature to
find different species of similar size where this type of
“conservation” exists. This and other factors point to a
delicate interrelationship which could be said to approach
parasitism. The present study is a description of some of
the behavioral aspects of this association.
METHODS ano MATERIALS
Observations and experiments, using SCUBA gear, were
made off the seaward side of the United States Coast
Guard Breakwater (hereafter referred to as the ‘Break-
water’) at the southwest end of Cannery Row, Monterey,
California, from 1965 to 1969, at depths of 50 to 60 feet.
A rectangular quadrant area of 3m < 33m _ was
marked off on the flat mud bottom running parallel to
the base of the Breakwater. The protected edge of the
study area was bordered by large boulders which form the
Breakwater. The study area contained 193 Cerianthus
individuals. Notes were kept on this population, which
remained stable, from October 5, 1968 to December 31,
1968. Cerianthus grow densely the length of the Break-
water, concentrated in a narrow strip close to its base.
An equal area directly to the seaward of the study area
supported only 11 animals.
Cerianthus were isolated on the bottom in cages made
of aluminum frame and screening, held in place by alu-
minum stakes pounded into the soft substrate.
Specimens of Dendronotus iris were of 2 distinct color
phases. Most had salmon-red bodies, and cerata and
branchial processes with orange ends, the tips of which
were white. In others the bodies were much lighter in
color, light milky purple with light orange processes,
tipped with white. The cerata and branchial processes
of both color phases sometimes had smaller purple side
branches. The largest specimen was 20cm long. There
seemed to be no relationship between color and size.
OBSERVATIONS
Feeding
Dendronotus iris feeds on Cerianthus tentacles. Over
a period of 2 months, 2 to 6 nudibranchs were found on
each dive in the study area. These were generally in
various attitudes of feeding. Because the average Ceri-
anthus tube projects above the substrate about 15cm
(larger specimens to 21 cm), the nudibranch must climb
the tube to reach the tentacles. Two methods of feeding
were recorded.
The first approach is used mainly by smaller (5 cm or
less) Dendronotus iris. The nudibranch climbs the tube
and slowly enters the rim of marginal tentacles. At this
point Cerianthus reacts by:
a. retracting swiftly and directly down its tube, D. iris
often climbing into the tube after the tentacles, or
b. allowing the nudibranch to crawl in among its partially
withdrawn tentacles and feed (Plate 55, Figure 1), or
Page 384
c. curling the ends of its marginal tentacles into tightly
coiled “pigtails,” in which case the nudibranch
crawls back down the outside of the tube and
leaves. This is uncommon and has only been seen
twice (Plate 57, Figures 5 and 6).
The second approach, in which the nudibranch is
pulled into the Cerianthus tube, is common among larger
nudibranchs. The nudibranch crawls up the outside of
the tube to within 35 mm of the lip. When one of its
spreading crown papillae, which form branched sheaths
surrounding retractile rhinophores, touches a tentacle,
an immediate and obvious stimulation of the nudibranch
takes place. The nudibranch raises the front part of its
body from the surface of the tube, arching the head back
as though peering up at the tentacles spread out above.
At the same time the mouth is thrust out baring large,
fleshy lips, and elongating the otherwise rather thick and
snubbed anterior end. Thus extended, the nudibranch
makes a sudden upward thrust with the anterior part of
its body, almost always catching one or more of the
marginal tentacles in its mouth (Plate 55, Figure 2).
The lips of D. iris form a vertical groove in which the
tentacle is aligned. As well as aligning the tentacle for
the jaws to close on, the lips seem to orient the nudi-
branch in relation to the position of Cerianthus. Some-
times after brief lip contact with the distal end of a
tentacle, the strike is made at the tentacle base. Dendro-
notus iris then gives a violent tug backwards, causing
Cerianthus to retreat quickly into its tube. The nudi-
branch hangs onto the tentacles with its jaws. In most
instances the force of the retreating tentacle(s) pulls the
nudibranch from its hold on the outside of the tube over
the lip, head first, down into the interior where it dis-
appears completely from sight (Plate 56, Figure 3). The
nudibranch stays inside the tube to complete feeding.
Time spent inside tubes varied from 20 minutes to over
24 hours.
Large Dendronotus iris sometimes miss in the attempt
to bite Cerianthus tentacles. The Cerianthus retract and
the top of the tube often collapses by folding, or the end
pinches closed. If this happens, most D. iris do not, or can
not, enter. If the tube remains open, the nudibranch
climbs in and feeds. To isolate the effect of Cerianthus
tentacles on D. iris, marginal tentacles were cut from live
THE VELIGER
Vol. 12; No. 4
Cerianthus. Three or 4 of these tentacles were held
with a test-tube holder so that the distal ends were free.
A D. iris was found crawling across the substrate (not on
Cerianthus). Cerata touched by anemone tentacles showed
no reaction. The tentacles were touched to the crown
papillae. As soon as contact was made, the Dendronotus
began extruding the mouth area (as described above) and
arched back the head exactly as it does when on a Ceri-
anthus tube. It extended its anterior end and made a
lunge forward, biting with its jaws as though biting at
tentacles.
A Dendronotus iris was placed on the tube of a retracted
Cerianthus. No tentacles projected from the tube. When
the crown papillae of D. iris were touched briefly with the
testing tentacles, D. iris reacted exactly as above, this time
making lunges toward the top of the tube, striking several
times with its jaws.
During one observation period 3 Dendronotus iris struck
at, but did not successfully hold onto Cerzanthus tentacles.
All 3 Cerianthus retracted and the nudibranchs left. Two
of the Cerianthus came out again within 2 hours. The
third remained within its tube for over 8 hours. None of
the 3 had its tentacles damaged by the D. iris encounter.
Sightings of such retractions and the re-emergence of Ceri-
anthus varied from 1 hour and 20 minutes to over 84
hours.
Quiescent Posture
Dendronotus iris is sometimes seen in a “sleeping” or
quiescent posture. In this posture a nudibranch clings to
the outside of a tube near the top, attached by the poste-
rior end of its foot. The anterior part of the foot is
unattached, and the back is arched. The head is held in
an upright position, and the nudibranch does not move,
except to sway back and forth with the currents. When
taken off the tube in this state, a nudibranch took a few
seconds to become active again. One D. iris maintained
the quiescent posture for over 8 hours.
Whether the quiescent posture represents a digestive
period, a recovery period, or otherwise, I cannot say.
Some nudibranchs crawl off the Cerianthus immediately
after feeding, some remain on the tube to lay eggs, and
others assume the quiescent posture.
Explanation of Plate 55
Figure 1: Dendronotus iris feeding among extended Ceranthus sp.
tentacles.
Figure 2: Dendronotus iris on Cerianthus sp. tube. Dendronotus iris
has just made a sudden upward thrust, baring large fleshy lips
which make contact with a Cerianthus sp. tentacle.
Tue VELIGER, Vol. 12, No. 4 [WopBeER] Plate 55
Figure 2
photographs by Don Woszer
Vol. 12; No. 4
THE VELIGER
Page 385
Escape
When Dendronotus iris is handled, it often swims by
means of heavy side-to-side gyrations of the body (Plate
56, Figure 4). This motion in light surge conditions, nor-
mal off the Breakwater, does not appear to orient the
nudibranch in a positive direction, but seems to be an
escape response to take it off the substrate. I did not
test this swimming reaction in the laboratory or under
controlled conditions. AGERsBorG (1922) states that swim-
ming is Dendronotus giganteus’ most common mode of
locomotion and assumes that it is pelagic in habit (D.
giganteus O’DonocHUuE, 1921 is synonymized with D.
iris by MAcFartanp, 1966). Dendronotus iris’ swimming
activities in Monterey seem restricted to escape reactions
only. Once off the bottom, D. iris is at the mercy of
water movement until it comes in contact with something
solid for a new foothold.
The active reef predator, Pycnopodia helianthoides
(BranpT, 1835), is often seen on the floor of the bay.
This sea star is able to excite Dendronotus iris into a
violent type of swimming escape response. In one obser-
vation, the anterior end of D. iris was inside the tube of
Cerianthus with only the posterior end visible. Pycnopo-
dia helianthoides was placed 15cm from the base of
Cerianthus. The sea star approached, one ray slowly ex-
tending towards the nudibranch. As soon as contact was
made, the nudibranch backed out of the Cerianthus tube,
released itself from the tube, and started gyrating. Water
currents carried it away. Twenty similar experiments with
P. helianthoides and different D. iris resulted in the same
immediate escape reaction.
The bat star, Patiria miniata (BRANDT, 1835), often
seen on Cerianthus tubes, will cause a slower and not so
violent swimming reaction of shorter duration. I have
never seen a sea star of any kind attack Dendronotus iris
in the field under natural conditions. It is of interest
that when a ray of Pycnopodia helianthoides is touched to
the Cerianthus, the sea star withdraws its ray.
Dendronotus iris is affected adversely when it comes in
contact with the anemone Tealia sp. which is found
among the Cerianthus beds. I have guided several swim-
ming D. iris onto Tealia. The tentacles immediately ad-
here to the D. iris and pull it toward the mouth where it
disappears into the gullet. Usually a slight escape response
is seen, but the response is weak and ineffective, as though
the nudibranch were partially paralyzed. When a single
Tealia tentacle was touched to D. iris, the nudibranch
turned aside and crawled in the other direction.
Reproduction
Dendronotus iris lays ringlets of white eggs on the
outside of Cerianthus tubes, attaching them at a point
high off the muddy bottom. This position may help in
avoiding predators, and additional protection may be
gained by proximity to the anthozoans’ tentacles. Eggs
have been seen during most of the months of the year
and at one time adults of all sizes are found. Copulation
takes place on the Cerianthus tube or on the mud bottom.
Copulation between the two color phases of D. iris is
common.
Locomotion
The expansive foot of Dendronotus iris is elongate,
flattened, and muscular. It is highly ciliated and has
good adhesive qualities (AGERsBorG, 1922). The foot is
adapted for travel on sand or mud bottom, and for
clinging to Cerianthus tubes in surge conditions.
Movements of crawling Dendronotus iris were traced.
Animals were seen to crawl past Cerianthus in what
seemed to be random directions. However, when crawling
away from Cerianthus beds, nudibranchs were seen to
reverse their direction. Whether or not a specific chemical
attractant or other signal is involved needs to be explored.
Tentacle Damage
The foraging habits of Dendronotus iris are such that
it goes from one Cerianthus to another. After being at-
tacked, the anthozoan stays within its tube. Cerianthus
near feeding D. iris are often retracted, as though having
been recently disturbed, even though 90 to 95% of the
Cerianthus outside the immediate area may be in the
expanded state. Repeated examinations of Cerianthus
tentacles after D. iris has fed on them, have shown only
minor damage with but 2 to 10 tentacles missing.
To get an indication of damage a single nudibranch
might do to a Cerianthus if both were confined, 2 nudi-
branchs over 16cm in length were kept in the field
without food for 48 hours. Cylindrical screen cages 48 cm
high by 30 cm diameter were placed over separate Ceri-
anthus. One “starved” nudibranch was placed in each of
the 2 containers. Within 15 minutes, both nudibranchs
had climbed the Cerianthus tubes, attacked the tentacles,
and were pulled inside the tubes, where they remained.
Four days later both nudibranchs were on the bottom of
the cages. When the cages were removed the nudibranchs
crawled away to feed on other Cerianthus. The 2 test
Page 386
THE VELIGER
Vol. 12; No. 4
Cerianthus remained retracted. Three days after the
cages were removed, only one test Cerianthus was out
and feeding. Only 20 to 30 tentacles appeared normal.
Others were eaten off at various lengths, more than half
left quite long. The top 6 cm of the tube of this Cerianthus
was tapered, white and smooth on its outside as though
newly secreted. Within one week it had darkened, thick-
ened and looked normal. The second Cerianthus was in
its tube, which was bent in an arc, and seemed lifeless.
Eleven days after the removal of the Dendronotus iris,
the first Certanthus tentacles had regenerated back to
normal, and the second had finally emerged from its
tube, the damage to its tentacles only slight.
Tentacle Curling
The tight end-curling of the marginal tentacles of Cerz-
anthus mentioned earlier has been observed on numerous
occasions. The exact function of this reaction is not known.
Usually, when disturbed, the tentacles are pulled inside
the tube with a single, swift motion. End-curling can be
induced artificially by lightly touching some of the ten-
tacles. As soon as contact is made all tentacles jerk in
unison about halfway in, then quiver and curl into the
typical “pigtail” shapes.
It is possible that tentacle end-curling is defensive,
that coiled tentacles expose a more concentrated mass of
nematocysts to predators. Whether a heavier concentra-
tion of nematocysts would affect Dendronotus iris is not
known. The curling could be a form of retreat, a method
of pulling in tentacles so they cover less area. Curling
could also be a reaction preparatory to withdrawal.
‘Tentacle end-curling has not previously been described,
and may not be a universal trait. Cerzanthus in Baja Cali-
fornia, Mexico, will curl the tentacles; however, the
anemones’ reflexes are much slower and show no prelim-
inary quivering. G. Robilliard reports seeing tentacle end-
curling in the Puget Sound area (personal communica-
tion). Dr. G. A. Horridge states that he has not observed
these curling reactions in the Naples species (personal
communication).
DISCUSSION
THompson (1962) remarks “nudibranchs preying on
organisms which form stable, abundant populations in
certain types of locality have annual life cycles; usually a
single breeding period each year.” A single breeding peri-
od coupled with a possible change of diet at maturity
would allow sedentary prey a respite from heavy cropping:
a period to grow and reproduce before the next mature
generation’s onslaught.
Dendronotus iris reproduces the year round. Nudi-
branchs of all sizes afford no relief to a standing crop of
Cerianthus, whose life span may cover a period of 10
to 40 years (Hyman, 1940, p. 632). While large nudi-
branchs and a constant harvest are a condition of exist-
ence, the continuing harvest of Cerianthus seems well
ordered and light, and the subtleties of the predator-prey
interrelationships contribute to what appears to be a well
balanced and stable ecosystem.
Dendronotus iris’ extensive branched crown papillae
are sensitive to Cerianthus tentacles. Stimulation of
these, regardless of the particular substrate the nudibranch
is on or its physical orientation, triggers a chain of
predictable responses which culminate in the biting action.
The fleshy mouth is well adapted for positioning tentacles
and the large prominent jaws for grasping and holding on.
Nematocyst cells in the Cerzanthus tentacles keep other
predators away. What actual effect they have on Dendro-
notus iris is uncertain. The curling response of the ten-
tacles and the fact that the tentacles are often curled
within the tube when the anemone is retracted are of
interest. Quick, complete withdrawal of tentacles might
be viewed as a way of escape, often causing the biting D.
iris to miss. This, combined with a closing or collapsing
tube, serves to frustrate attack. On the other hand, it
seems more than a drastic outcome for the anemone to
pull the nudibranch into its tube. Once the nudibranch
is in the tube, its feeding may be restricted. Perhaps a
nematocyst threshold, mucus emission, low oxygen level,
the nudibranchs’ limited digestive capacity for Ceran-
Explanation of Plate 56
Figure 3: Dendronotus iris in the process of being pulled into the
tube of the anthozoan, Cerianthus sp.
Figure 4: Dendronotus iris “swims” by means of heavy side to side
gyrations of the body. This appears to be an escape reaction.
Explanation of Plate 57
Figure 5: Dendronotus iris among Cerianthus sp. tentacles. Note
that the tentacles close to the anterior end of the nudibranch are in
the tightly coiled “pigtail” configuration.
Figure 6: Detail of the Cerianthus sp. tentacle end-curling.
THE VELIGER, Vol. 12, No. 4 [WopBerR] Plate 56
Figure 3
Figure 4
photographs by Don WosBER
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Vol. 12; No. 4
THE VELIGER
thus tentacles, or other factors terminate the feeding and
ultimately drive the nudibranch out. Data on how long
the nudibranch stays in the tube vary from 20 minutes to
over 24 hours. Regardless of the length of time spent in
the tube, extensive damage to Cerianthus has not been
seen in the field. The tube itself limits predation and
Cerianthus can exist long periods of time inside the tube
without coming out.
ACKNOWLEDGMENTS
The help and assistance of Dr. Robert D. Beeman, San
Francisco State College; Dustin D. Chivers, California
Academy of Sciences; Dr. Michael T: Ghiselin, University
of California; Dr. Cadet Hand, Bodega Marine Labora-
tory; Gordon A. Robilliard, University of Washington; and
Dr. Charles R. Stasek, Florida State University in the
preparation of this paper is gratefully acknowledged.
Page 387
LITERATURE CITED
Cooper, JAMES GRAHAM
1863. | On new or rare mollusca inhabiting the coast of Cali-
fornia. Proc. Calif. Acad. Nat. Sci. 3 (2): 56 - 60
Hyman, Lippy HENRIETTA
1940. The invertebrates. vol. 1: Protozoa through Ctenophora.
McGraw-Hill, New York, i- xiit+ 726 pp.; 221 figs.
KyerscHow-Acerssorc, H. P.
1922. Notes on the locomotion of the nudibranchiate mollusk
Dendronotus giganteus O’DoNoGHUE. Biol. Bull 42: 257
to 266
MacFar.anp, FRANK Mace
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
TuHompson, Tuomas E.
1962. Grazing and the life cycles of the British nudibranchs.
In: D. J. Crisp (ed.): A symposium of the British Ecological
Society, 1962. Blackwell Sci. Publ., Oxford, pp. 275 - 285
Page 388
THE VELIGER
Vol. 12; No. 4
Cargoa cupella, New Genus and New Species of Nudibranch
from Chesapeake Bay and the Generic Status
of Okenia MENKE, Idalia LEucKART and Idalla ORsTED '
ROSALIE M. VOGEL
AND
LEONARD P. SCHULTZ
University of Maryland, Chesapeake Biological Laboratory, Solomons, Maryland 20688
(5 Text figures)
Durinc A stupy of the literature concerning the identity
of a rare nudibranch collected in Chesapeake Bay, it was
observed that species in this relationship (Suborder Eu-
doridacea, Superfamily Suctoria, Family Goniodorididae)
were usually placed in the genus Jdalia LEucKaRT, 1828
or in Okenia MeENnKE, 1830. An attempt was made to
determine the valid generic taxon available for the species
in this group.
The authors gratefully acknowledge the drawings made
by Mrs. A. J. Mansueti and especially thank Mr. David
G. Cargo, who collected the holotype, and called the spe-
cimen to our attention. Also we appreciate the use of the
facilities of the Chesapeake Biological Laboratory, Solo-
mons, Maryland. The staff of the Division of Mollusks,
Smithsonian Institution, kindly let us use their library and
made specimens from their collections available to us.
To: Drs. David R. Franz, University of Connecticut;
Joseph Rosewater, Smithsonian Institution; Anne Hurst,
University of Reading, United Kingdom; and Mrs. Eve-
line Marcus, Sao Paulo, Brazil, we are pleased to acknow-
ledge their valuable suggestions upon reading this manu-
script before it was submitted for publication.
This research was supported in part by the Bureau of
Commercial Fisheries and the Department of Chesapeake
Bay Affairs, Contract no. 14-17-0007-959 under Public
Law no. 89-720 of the Jellyfish Act.
The nomenclatural history of this group is given below.
Only the more important references are cited in the syn-
onymy. ‘
‘ Contribution Number 4030
Cargoa VocEL & SCHULTZ, gen. nov.
Type species: Cargoa cupella VocEL & SCHULTZ, spec. nov.
We are pleased to name this new genus in honor of our
colleague, David G. Cargo, Chesapeake Biological Labor-
atory, Solomons, Maryland.
Okenia LeucKartT in Bronn, 1826, Ergebn. Reisen, vol. 1,
p. 329 (O. elegans LEUCKART, nomen nudum).
Okenia MENKE in IREDALE & O’DoNoGHUE, 1923, Proc.
Malacol. Soc. London, vol. 15, prt. 4, pp. 197, 217
(nomenclature, preoccupied in Diptera by Okenia
ZETTERSTEDT, 1838). — MENKE in Basa, 1937, Nov.
20, Journ. Dept. Agric. Kyushu Impl. Univ., vol. 5,
p. 295 (Type-species designated, Idalia elegans LEuc-
KART). — PruvoT-Fot, 1954, vol. 58, p. 308. — Mar-
cus, 1957, Journ. Linn. Soc. London, vol. 3, p. 436
(nomenclature). — Macnae, 1957, Trans. Roy. Soc.
South Africa, vol. 35, p. 368 (nomenclature) .
Idalia LeucKarT, 1828, Breves anim. quorund. descr. p.
15, fig. 2a, 2b. (Type-species, I. elegans LeucKarr,
p. 15; preoccupied by Jdalia HuEBNer, 1819 and
Idalia HuEBNER, 1825, in Lepidoptera, ref. copied
from Neave).— Menke, 1830, Synop. Method. Mol-
lusc. p. 10 “(Idalia Leuckart = Okenia Leuc-
KART)”. — Puippt, 1844, Fauna Moll., p. 76 (genus
described; 6 species listed). — ForBEs & HANLEy,
1853, History Brit. Moll., vol. 3, p. 578. — ALDER &
Hancock, 1855, Monogr. Brit. Moll., genus 8, fami-
ly 1, (genus divided in two sections. Sec. 1 is desig-
Vol. 12; No. 4
THE VELIGER
Page 389
nated “Type, J. elegans,” plt. 27, figs. 1-5; also see
app. p. xviii; Sec. 2, “Type, I. aspersa,” plt. 26, figs.
1-10, which Bercu, 1881 placed in Jdaliella
BercH ). — Bercu, 1881, p. 142, subgenus on p. 144
(Idalia). — Bercu, 1883, Beitrage Monog. Polycera-
den, Wien, p. 41, (subgenus Idalia, p. 42). — Exiot,
1910, Brit. Nudibranch. Moll., pt. VIII (suppl.), p.
158 (continued to divide genus into 2 subgenera fol-
lowing Bercu, 1881).
Idalina Norman, 1890, Ann. Mag. Nat. Hist., ser. 6, vol.
6, p. 74 (replaces Idalia LeucKart, 1828; Idalina
NorMaN preoccupied in Protozoa by Idalina Mu-
NIER-CHALMAS & SCHLUMBERGER, 1884, ref. copied
from NEAvE) ;— IREDALE & O’DoNnocHUuE, 1923, Proc.
Malacol. Soc. London, vol. 15, prt. 4, p. 197, (“Ida-
lina is altered to Okenia’’).
Diagnosis: The new genus Cargoa is distinguished from
other genera in the family Goniodorididae by the combi-
nation of the following characters: Tentacles clavate or
linear; leaves present on non-retractile rhinophores; non-
retractile gills present; one row of papillae present on
each side on a pallial ridge; one or more dorsal papillae
present on back. Additional characters of the type-species
are those of the new genus.
Okenia LeucKart in Bronn (1826) is anomen nudum
because O. elegans LrEucKart (1826) was without de-
scription. Only a name was listed.
MeENKE (1830) listed Idalia LeuckART — Okenia
Leucxkart, also without a description. Okenia here is a
synonym of Idalia. However, it is not available as of
MeENKE (1830) because in the Code, Art. 10d, “A name
first published as a synonym is not thereby made avail-
able.” The Code, Art. 16b, ii, also covers this problem. We
quote Art. 16b, “The following are not ‘indications’ in the
meaning of this chapter: (11) citation of a name in syn-
onymy [Art. 11d]. We conclude therefore that Okenia
MeEnKE (1830) is not available. Okenia MENKE next
appeared in IREDALE & O’DonocHuE (1923) and in
Baza (1937), the latter designated J. elegans LEUCKART
as type-species. However, if Okenia MENKE in IREDALE &
O’DonocHuE (1923) or in Basa (1937) is considered
valid, it is preoccupied in Diptera by Okenia ZETTER-
STEDT, 1838.
Norman in 1890 proposed Idalina to replace Idalia
Leucxkart, 1828, but Idalina Norman, 1890 is preoccu-
pied by Idalina Munter-CHatmaAs & SCHLUMBERGER,
1884.
Leuckart in 1828 proposed Idalia elegans and gave a
description, but Idalia is preoccupied in Lepidoptera by
Idalia HueBNer, 1819. Bercu in 1881 accepted Idalia
Leucxart but divided it into two subgenera, naming one
of them Idaliella with I. aspersa (ALDER & Hancock) as
the type-species. The latter species differs from Idalia ele-
gans by lacking a mid-dorsal papilla, whereas I. elegans has
one or more papillae on the back. This difference is great
enough to indicate two distinct genera.
We conclude that the group of species usually referred
by authors to Idalia or Okenia is without an available
generic name. Thus we now propose Cargoa, new genus
with the type-species, Cargoa cupella, new species. This
new genus is not a replacement name for either Jdalia or
Okenia.
We believe that the following species should be assigned
to the new genus, Cargoa:
C. elegans (Leuckart, 1828) (type-locality, Mediterra-
nean).
C. cirrigera (Puiwiprr, 1839) (type-locality, Sicily).
C. leachu (ALDER & Hancock, 1854) (type-locality, Tor-
bay, Whitburn, Durham, Hebrides).
C. tentaculata (Stimpson, 1855) (type-locality, China).
C. mediterranea (IHERING, 1886) (type-localtiy, near
Naples, Mediterranean).
C. plebia (BercuH, 1902) (type-locality, coast of Lem
Ngob, Siam).
C. dautzenbergi (VaysstzrE, 1919) (type-locality, Gulf of
Marseille, Mediterranean).
C. vancouverensis (O’DonocHuE, 1921) (type-locality,
Nanaimo, Vancouver Island, British Columbia,
Rosepit, north end of Queen Charlotte Islands).
. distincta (BaBA, 1940) (type-locality, Asamusi, Japan).
. japonica (Basa, 1949) (type-locality, off Sajima, Sa-
gami Bay, Japan).
. echinata (Basa, 1949) (type-locality, off Sajima, Sa-
gami Bay, Japan).
. evelinae (Marcus, 1957) (type-locality, Ubatuba and
Ilhabela, Brazil; Florida).
. impexa (Marcus, 1957) (type-locality, Island of Sao
Sebastiao, and Ubatuba, Brazil).
. opunta (Basa, 1960) (type-locality, Tannowa, Osaka
Bay, Japan).
. plana (Basa, 1960) (type-locality, Toba, Japan, and
Kada, Osaka Bay, Japan).
. babai (Hamatani, 1961) (type-locality, Tannowa,
Osaka Bay, Japan).
C. angelensis (LANCE, 1966) (type-locality, Bahia de Los
Angeles, Estado de Baja California, Mexico).
C. pellucida (Burn, 1967) (type-locality, Sidney Harbour,
New South Wales).
C. mija (Burn, 1967) (type-locality, Point Danger, Tor-
quay, Victoria).
C. sapelona (Marcus & Marcus, 1967) (type-locality,
Sapelo Island, Georgia) .
Ye). De Se SS) Ss SS)
Page 390
Cargoa cupella VocEL & SCHULTZ, spec. nov.
(Figures 1 to 5)
Holotype, USNM 679396 (deposited in Division of Mol-
lusks, U. S. National Museum). A specimen 2 mm (crawl-
ing length) long, collected at Aberdeen Rock, York River,
Virginia, by David G. Cargo on November 1, 1968.
Paratype, radula from a specimen, USNM 679397, bear-
ing same data as holotype.
Description based on the living and crawling specimens:
Cargoa cupella has a narrow and fairly long body, its
height about 4.6 and its width 6.2 times in total length.
The animal is compressed laterally so that the height of the
animal in the cardiac region is higher than it is broad.
The foot is not as wide as the body. The anterior end of
the foot is slightly bulbous and rounded.
The highest point in the dorsal profile is at the base
of the mid-dorsal papilla. The heart is dorsal but does not
form a protuberance noticeable on the surface. One row
of 7 lateral papillae occurs on each side of the back. ‘These
papillae are supported by spicules and are connected bas-
ally by a thin mantle or membranous veil which forms the
pallial ridge. The edge of the mantle is a smooth scalloped
line between the papillae. The first pair of papillae is
located just laterally to the base of the rhinophores and
the last pair which is clavate and joined at the base, is
located just postero-laterally to the tips of the 4 branchial
plumes. A single elongate clavate papilla is located in the
mid-dorsal region just posterior to the heart. Posterior to
THE VELIGER
Vol. 12; No. 4
the single mid-dorsal papilla are located 4 pinnate, bran-
chial plumes. These can be flattened against the body but
cannot be retracted. The rhinophores have blunt ends and
are situated in the head region. They are long, slender,
non-retractile structures although they can be pressed
downward against the body of the animal. The cup-
shaped leaves facing distally are located about 2 the way
out the length of each rhinophore.
There is a pair of tentacles anterior to the rhinophores.
The tentacles are about half the length of the rhinophores
and are cylindrical, with rounded tips. The tentacles are
not clavate in shape. They usually slant upwards and for-
ward, but on occasion have been observed stretching for-
ward and downward until they touch the substrate on
which the animal was crawling.
The tail, or the posterior end of the foot, is broad and
flat, and widens in the proximal 4, then tapers distally.
The head is not divided as such from the body. It is thick
and slopes slightly to the upper lip. The mouth is subter-
minal, and is surrounded by fleshy lips. The eyes are vis-
ible dorsally in the head but are more easily seen from
the side since they are deeply set in the translucent body
tissues. They appear as large black dots.
The anal opening is just posterior to the branchial
plumes in the mid-dorsal position. The reproductive open-
ings are on the right side below the second papilla. The
penis is armed with barbs.
The animal is of a whitish coloration with white spots
concentrated in the papillae and rhinophores. There are
dark brown or black spots on the back, anterior to the mid-
dorsal papilla. These may be rather sparse or quite con-
centrated to form a definite pattern between the rhino-
phores and almost to the branchia, sometimes not ex-
Figure 1
Cargoa cupella VocEL « SCHULTZ, spec. nov.
drawing by Alice Jane Mansueti
Lateral view of holotype, USNM 679396
b — branchia
s — spicules
a — anus
le — leaves
t — tentacle
p — papillae
Vol. 12; No. 4
THE VELIGER
Page 391
tending past the base of the mid-dorsal papilla. The
longest branchia (middle two) are 4 the length of the
rhinophores. The rhinophores are approximately 0.7 mm
long.
The radula is composed of approximately 10 rows of
4 teeth each. The dental formula is 1:1:0:1:1. The teeth are
pointed posteriorly. On the inner edge of the inner lateral
teeth are 9 denticles.
Four specimens of this species were found crawling on
an oyster shell containing Chrysaora quinquecirrha polyps
and other common fouling organisms. The shell was col-
Figure 2
Cargoa cupella VocEL & SCHULTZ, spec. nov.
drawing by Alice Jane Mansueti
Dorsal view of holotype, USNM 679396
r — reproductive openings on right side
beneath second lateral papilla (not shown in drawing)
lected at Aberdeen Rock, York River, Virginia, on 1 No-
vember 1968. The salinity was 20% . These nudibranchs
were kept alive in aquaria for observation of their behavior
and to have drawings made of them. During a 10-day
period 2 were lost before they could be dissected, one was
preserved (holotype), and the radula was dissected from
the other one. The 2 specimens that disappeared showed
little or no variability from the holotype.
One animal laid an egg mass that was attached along
its lower part to the side of the glass bowl. It was 1 mm in
length, oval in shape, and contained approximately 40
white eggs, that developed to beyond the 32-cell stage and
then died.
1.0mm
Figure 3
Cargoa cupella VocEL & SCHULTZ, spec. nov.
drawing by Alice Jane Mansueti
Ventral view of holotype, USNM 679396
Page 392
THE VELIGER
Vol. 12; No. 4
0.005 mm
Figure 4
Cargoa cupella VocEL &« SCHULTZ, spec. nov.
drawing by Alice Jane Mansueti
Radula of paratype, USNM 679397
1 - lateral tooth m — marginal tooth
Figure 5
Cargoa cupella VocEL & SCHULTZ, spec. nov.
drawing by Alice Jane Mansueti
Egg mass
The animals were exposed to Chrysaora quinquecirrha
polyps, several sponges and hydroids, in an effort to see
what they ate. None of these items were seen taken by the
nudibranchs.
Remarks:
This new species differs from the other members of the
genus by having 3 to 5 cup-shaped leaves on the rhino-
phores, whereas Cargoa evelinae and C. sapelona have 6;
C. plebia has 25; C. vancouverensis has 28 - 30; C. im-
pexa has 8 or 9; C. pellucida has 7 - 12; C. babai and C.
mija have 4 - 6. The species — C. elegans, C. cirrigera, C.
leachu, C. mediterranea, C. distincta, C. japonica, and C.
opunta, all have perfoliate rhinophores, whereas C. cu-
pella has 3 to 5 cup-shaped leaves on the rhinophores.
Both Cargoa cupella and C. tentaculata possess 4 gills,
one dorsal papilla, and one pair of anterior tentacles.
However, C. tentaculata differs by having large, clavate
rhinophores in contrast to the slender, cylindrical rhino-
phores bearing 3 to 5 leaves for C. cupella. Cargoa tenta-
culata is wine yellow with brownish spots along each side
of the flake-white median line. It has chestnut brown rhi-
nophores and gills. Cargoa cupella, in contrast, is whitish
with bright white spots concentrated in the rhinophores
and papillae. It also has dark brown or black spots dorsal-
ly from the rhinophores almost to the gills.
While this paper was in the hands of the editor, 2 ad-
ditional specimens of Cargoa cupella were collected No-
vember 4, 1969, one from the Manokin River, and another
from Deal Island; this latter specimen was dead and
deteriorating. The former, 4mm in length, had 5 cup-
shaped leaves on each rhinophore. This indicates that the
cup-shaped leaves of C. cupella may increase in number
with increase in length. Therefore, among the 20 other
species referred to the genus Cargoa, the number of
leaves 3 to 6 on C. evelinae, C. mija, and C. sapelona, over-
lap with the number 3 to 5 as found on C. cupella. The
latter differs by having only one large club-shaped papilla
on the mid-dorsal line which distinguishes it from C.
evelinae, C. mija, and C. sapelona, which have 4, 9 and
5, respectively.
This new species is named cupella (gender, feminine)
in reference to the small, cup-shaped leaves on the rhino-
phores.
Idalla Orstep, 1844
During our investigation of the nomenclatural history
of Idalia and Okenia, we noted that Orstep (1844, p. 73)
listed Idalla caudata Orstepv from Kullen, Sweden. In a
footnote, which we translate, he described the species as
follows:
Body oblong, much higher than wide, golden, tail
slender and curved upward, 3 simple lappets on both
sides dorsally, 6 tentacular appendages, whereby 2
longer are double, 8 anal gills, length 4-5 mm, width
2mm.
It should be noted that he does not mention a mid-
dorsal papilla as found in Cargoa. The other characters of
Idalla caudata indicate that it should be referred to the
group of species included in Idaliella Bercu, 1881. In
reviewing the nomenclatural history of Jdaliella Brercu
(1881, p. 145), we found that [dalla Orstep has priority
over the usually recognized genus Idaliella with Idalia
aspersa ALDER & Hancock, 1855 as the type-species.
The following species, among others are referred to this
genus: I. caudata Orstev; 1. aspersa (ALDER & HANcocK);
Vol. 12; No. 4
THE VELIGER
Page 393
I. pulchella (ALDER & Hancock); I. fusca (ODHNER) ; I.
inaequalis (Forses & Hanwey); I. quadricornis (Mon-
TAGU) ; I. amoenula (BERGH) ; I. barnardi (BaBA).
Some authors have placed Euplocamus Puteri, 1836
(type-species, EF. croceus Puitippi1; preoccupied by Eu-
plocamus LATREILLE, 1809 in Lepidoptera) with Idaliella.
We do not think the two genera are the same because E.
croceus does not belong zoologically with the species
herein referred to Jdalla.
LITERATURE CITED
ALDER, JosHUA & ALBANY Hancock
1845 - 1855. A monograph of British Mollusca.
prts. 1-7 [no pagination]
Basa, K1KuTARO
1937. | Opisthobranchia of Japan (II). Journ. Dept. Agr.,
Kyushu Imp. Univ., Fukuoka 5 (7): 289 - 344; plts. 1, 2
(20 November 1937)
1940. Some additions to the nudibranch fauna of the northern
part of Japan. Bull. Biogeogr. Soc. Tokyo 10 (6): 103 - 111;
fig. 1-10 (March 1940)
1949. Opisthobranchia of Sagami Bay, collected by his Majesty
the Emperor of Japan. Tokyo. 104 pp.; 50 plts.; 7 text figs.
Biol. Lab. 8 (1) :79- 85; plts. 7, 8
Ray Soc.,
(November 1949)
1960. The genera Okenia, Goniodoridella and Goniodoris from
Japan (Nudibranchia - Goniodorididae) . Publ. Seto Mar.
Biol. Lab. 8 (1): 79-85; plts. 7, 8 (30 May 1960)
Bercu, Lupwic SopHus RupDOLF
1881. Uber die Gattung Idalia LrucKarr.
gesch. 47 Jahrg. 1: 140 - 181; plts. 6-8
1883. Beitrage zu einer Monographie der Polyceraden. Prt. 33:
1 - 48; plts. 6- 10 (7 March 1883)
Bronn, HeEinricH GEORG
1826. | Ergebnisse meiner naturhistorisch Gconomischen Reisen.
1: 329
Burn, RoBert
1967. Descriptions of two new species of Okenia (Nudibran-
chia, Doridacea) from southeastern Australia. Proc. Roy.
Zool. Soc. New South Wales: 52-58; figs. 1-5; plts. 10, 11
(24 February 1967)
Arch. Natur-
Exiot, Cartes N. E.
1910. The British nudibranchiate Mollusca.
Soc. 8: 1-198; plts. 1-8
Forses, Epwarp & SYLVANUS HaNLEY
1853. History of British Mollusca.
Hamatanl1, Iwao
1961. Preliminary account of a new species of Okenia from
Osaka Bay, Japan (Nudibranchia-Goniodorididae) . Publ.
Seto Mar. Biol. Lab. 9 (2): 363 - 365; fig. 1
(20 December 1961)
International Commission on Zoological Nomenclature
1961. International code of zoological nomenclature. pp. 1-176
Suppl. Ray
London, 3: 1 - 161
IREDALE, ToM & CHARLES HENRY O’DoONOGHUE
1923. List of British nudibranchiate mollusca. Proc. Malac.
Soc. London 15: 195-200 (March) ; 201 - 233 (June, 1923)
Lance, JAMES RoBERT
1966. New distributional records of some northeastern Pacific
Opisthobranchiata (Mollusca:Gastropoda) with descriptions
of two new species. The Veliger 9(1): 69-81; figs.
1-12 (1 July 1966)
LreuckarT — _ see BRONN
LEUCKART, FRIEDRICH SIGISMUND
1828. Breves animalium quorundam maxima ex parte marin-
orum descriptiones.
Macnae, WILLIAM
1957. The families Polyceridae and Goniodorididae (Mollusca,
Nudibranchiata) in southern Africa. Trans. Roy. Soc. S.
Africa 25 (4): 341 - 372; figs. 1 - 23; plts. 17, 18
Marcus, ERNsT
1957. On Opisthobranchia from Brazil II.
Soc. London 43: 309 - 486; figs. 1 - 246
Marcus, EvELINE Du Bois RayMonpb & ERNST Marcus
1967. | Some opisthobranchs from Sapelo Island, Georgia, U. S.
A. Malacologia 6 (1-2): 199 - 222; figs. 1 - 17
(December 1967)
pp. 1 - 24; figs. 1-7
Journ. Linn.
MEeENKE, Kart THEODORO
1830. Synopsis methodica molluscorum. pp. 1 - 169
NEAVE, SHEFFIELD AIREY
1939-1940. Nomenclator zoologicus. Zool. Soc. London;
vol. 2: 1 - 1025; vol. 3: 1 - 1065
Opuner, Nirs HjALMar
1939. | Opisthobranchiate mollusca from the western and north-
Kgl. Norska Vidensk. Selsk. Skr.
(20 February 1939)
ern coasts of Norway.
1: 1 - 93; 59 text figs.
O’DonocGHuE, CHARLES HENRY
1921. | Nudibranchiate molluscs from the Vancouver Island
region. ‘Trans. Roy. Canad. Inst. 13 (1): 147 - 209; plts. 1-4
(February 1921)
OrstTED, ANDERS SANDGE
1844. De regionibus marinis elementa topographie historico-
naturalis Freti Oresund. pp. 1 - 88; plts. 1, 2
Puiippl, RuDOLFUS AMANDUS
1844. Fauna molluscorum.
ton, pp. 1 - 304; plts. 13 - 28
Pruvot-Fo., ALICE
1954. | Mollusques opisthobranches.
to 460; 1 plt.; 173 text figs.
plts. 1-4
Stimpson, WILLIAM
1855. Descriptions of some new marine invertebrates from the
Chinese and Japanese Seas. Proc. Acad. Nat. Sci. Phila-
delphia 7 (10): 375 - 384
VAYSSIERE, ALBERT JEAN BapTisTE Marie
1919. Recherches zoologiques et anatomiques sur les mollusques
opisthobranches du Golfe de Marseille. 2nd suppl. 17:
53 - 92
ZETTERSTEDT, J. Wm.
1838. Insecta Lapponica descripta
Halis Sar. Sumt. Eduardi An-
Faune de France 58: 1
[reference copied]
Page 394
THE VELIGER
Vol. 12; No. 4
Malacological Applications of Scanning Electron Microscopy
I. Introduction and Shell Surface Features
BY
ALAN SOLEM
Department of Zoology, Field Museum of Natural History
Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605
(Plates 58 to 60)
OBSERVATION AND ILLUSTRATION of shell features in micro-
mollusks have presented nearly insoluble problems to gen-
erations of malacologists. Using an optical microscope,
details cannot be seen at very low magnifications, while at
high magnification the depth of field is extremely shallow
and there is not enough light available to distinguish
finer structures. Photographic efforts generally are even
less successful. The low resolving power of camera lenses,
their very shallow depth of field, lighting problems, and
the graininess of the film emulsion result either in minute
images with little detail or large pictures with some areas
obviously out of focus and details in other regions obscured
by dark shadows. Various ingenious lighting techniques
can reduce some of these difficulties (BLAKER, 1961), but
nothing can solve the optical depth of field limitations.
Use of a “camera lucida” to prepare outline drawings of
small shells or to establish proportions for an artist to make
either a pen and ink or tone drawing has been a partially
effective alternative. The Wild M-5 stereoscopic dissecting
microscope with camera lucida, for example, is a superb
instrument and very easy to use. Considerable time in re-
constructing out of focus areas is required, the cost of
illustration is high, and fine details are completely absent.
In the last three years, commercial versions of the
scanning electron microscope have been installed in a
number of universities and industrial laboratories. Al-
though there is as yet little published work on mollusks
resulting from use of this instrument, already it is evident
that within a very few years the scanning electron micro-
scope (hereafter SEM) will be the routine working tool
for both examination and illustration of small mollusks.
Its impact on pollen studies and entomology already has
been dramatic. Even when renting SEM time at commer-
cial rates, the cost of illustrations is competitive with good
quality art work. Both the information content and quality
of the SEM photographs are far superior to optical photo-
graphs or drawings.
In the past 18 months I have been able to spend about
25 hours studying shell sculpture and radulae with a
Cambridge Mark I Stereoscan microscope at Alpha Re-
search and Development, Inc. of Blue Island, Illinois. Mr.
John A. Brown, electron microscopist, and his assistants
have been of invaluable help in interpreting puzzling fea-
tures. Equally important, their skill and efficiency in oper-
ation of the SEM has enabled completion of an amazing
number of photographs and opened up whole new vistas
for research. An equal debt is owed to the National Science
Foundation who permitted illustration funds from NSF
GB-6779 to be used in making the photographs repro-
duced here and to the administration of Field Museum
for providing additional funds for exploratory studies. Mr.
Fred Huysmans of the Field Museum photography depart-
ment took infinite patience in coaxing maximum informa-
tion from the polaroid negatives.
The present report is intended as an introduction to the
capabilities of the SEM and its applicability to the study
and illustration of shell surface features. A second report
will cover data on radulae. Papers by WisE & Hay (1968a,
1968b), Taytor, KENNEDY & Hatt (1969), ERBEn,
Frajs & SmeHL (1968), and Wise (1969) have been
concerned with cross-sectional structures of molluscan
shells. THompson & Hinton (1968) and Sotem (1969)
have published popular notes on shell sculpture studies. An
extensive bibliography of scanning electron micrography
is contained in the same volume as Wise (1969). Hay &
SANDBERG (1967) give an excellent introduction to the
subject. Most of my own comments concerning technical
specifications are paraphrased from these sources or from
information provided by John Brown of Alpha Research.
Vol. 12; No. 4
OPTICAL versus SEM VIEWING
Advantages of the SEM over optical equipment lie in a
50 to 500 increase in depth of field at a given magnifica-
tion, 124 to 100 increase in resolving power, a magni-
fication range of 20 to 50000 requiring clicks of a
single dial, and the ability to rotate, tilt, and turn the spe-
cimen being examined while keeping it continuously in
view. A comparison of the best quality stereoscopic dis-
secting microscope, compound light microscope, and SEM
capabilities is given in Table I. This is modified from Hay
& SANDBERG (1967, p. 410) with some additional data.
Optical systems have a depth of field that may be only
one-twentieth the field of vision width, but the SEM has
a depth of field that can reach five times the field of vision
width. The significance of this in illustrating globular ob-
jects such as shells cannot be overemphasized. Using the
lowest magnification (20>) of the Cambridge Mark Ila
Stereoscan instrument at standard working distance, for
example, any cylindrical shell whose length is less than
7mm can be photographed in a single exposure and the
image fills a 34x34 inch negative. Specimens of up to 12
mm in length can be photographed in two shots with a
montage print producing one continuous image for
reproduction. All portions of such a shell will be in focus.
The lighting effect will approximate that of natural sun-
light on a cloudy day. Details will be visible in all areas
and proper contrast control will prevent heavy shadow
effects. Heliciform shells of 5 mm diameter can be photo-
THE VELIGER
Page 395
graphed in one shot. Even all parts of the apertural view
will be completely in focus and details can be seen far
inside the aperture. Montage photographs of 5 - 10mm
heliciform specimens would be possible with 3 to 4 ex-
posures. Since the specimen position can be altered under
fine control, after the first photograph is taken, tilting of
a few degrees and taking a second photograph will yield a
stereo pair. Standard photogrammetric equipment such
as the Wild ST-4 Mirror Stereoscope with parallax bar
will enable relative depth measurements of surface topo-
graphy and three-dimensional viewing of the specimen.
At the lowest magnifications, the SEM can produce
whole specimen illustrations of 5 to 7 mm shells. A magni-
fication dial permits parfocal switching between 20, 50,
100, 300, 1 000, 3.000, 10 000, 30 000, and 50 000%: in
order to inspect structural details. Figures 3 to 11 present
part of such a series. At each step the specimen can be
moved and adjusted, changing the angle of view or
shifting to a different area. It is visible at all times unless
moved out of the scanning field by too enthusiastic
spinning of controls! If “lost”, returning to a lower magni-
fication will relocate and enable retrieval of the strayed
item.
SEM OPERATION
A series of electromagnetic lenses in a vacuum chamber
focus a beam of electrons into an electron probe of 50 to
Table 1
Comparative Instrument Performance
Stereoscopic Compound Scanning
Binocular Light Electron
Microscope Microscope Microscope
Resolution 20000 A 2500 A 200 A
Field of view at:
20 X 12.5 mm circle not normally used 5 X 5mm
9.4 mm square
50 X 4mm circle 2.1mm circle 2X 2mm
3 mm square
500 — 0.21 mm circle 0.2 X 0.2 mm
5 000 X — — 0.02 X 0.02 mm
50 000 X = — 0.002 X 0.002 mm
Depth of field at:
20 X 0.5 mm
50 X 0.2 mm
500 X —-
5 000 X —
50 000 x ==
not normally used + 20mm
0.02 mm 10mm
0.002 mm 1mm
— 0.1 mm
— 0.01 mm
Page 396
THE VELIGER
Vol. 12; No. 4
75A diameter. When this beam hits a surface, a few
electrons are reflected, but a much greater number of
low energy secondary electrons are emitted from the sur-
face. These electrons are caught by a detector, the signal
amplified several thousand times, and displayed as a spot
on a long persistence phosphor cathode ray tube. Image
brightness is determined by the intensity of secondary
electron emission, with a low emission rate producing a
dim spot and a high emission rate producing a bright spot.
Use of a sawtooth generator scans the focused electron
beam across the specimen. By making a quick scan, it is
possible to build and maintain a complete visual image
on the long persistence phosphor cathode ray tube. It is
this electronic image that one watches. Movements of
the specimen produce a few seconds of distorted image
until the previous scan spots fade and a new image builds
up.
Producing a photograph requires use of a second cath-
ode ray tube. In 100 seconds a single slow scan sweep is
made onto a low persistence phosphor cathode ray tube.
During this sweep the tube image is exposed to Polaroid
Type 55 positive-negative film. After 20 seconds develop-
ing time, a Polaroid black and white print plus a negative
are available for inspection. If brightness, contrast, or ex-
posure was miscalculated, then a second picture can be
taken immediately. About 5 minutes are required for
each exposure, including control adjustments after magni-
fication and specimen positioning have been selected.
About 10 publishable exposures can be made per hour.
Since the image was scanned onto the cathode ray tube,
attempts to enlarge a photograph more than 3 or 4 soon
reveal the raster lines familiar to all TV watchers. En-
largement of the upper right hand corner in Figure 1 to
full picture size (Figure 2) illustrates this problem. Huge
enlargements can be made by “fuzzing” the enlarger focus,
but this results in significant detail loss.
SPECIMEN PREPARATION
AND SIZE LIMITS
For dissipation of any electron charge that builds up
on non-conductive surfaces, a conductive surface is de-
sirable. Metallic substances can be examined directly, but
biological specimens and fossils either must be given a
conductive coating or else have the electron beam reduced
to between 1 to 3 kV. Great loss of resolution in the
image results from the latter option. Gold, palladium,
aluminum, platinum and carbon are commonly used coat-
ing materials. For effective viewing, a clean surface is
essential.
The specimens illustrated in this report were first cleaned
of most dirt and incrustations by soaking overnight in
water and then immersion for a few seconds into a water-
filled tank of an ultrasonic cleaner. They were attached
to a SEM stub with ordinary rubber cement and then
placed in a vacuum evaporator. After pressure had been
reduced to 10° torr, a 200 - 300 A layer of gold or alumin-
um was vaporized electrically in a tungsten filament
while the specimen was rotated slowly beneath the fila-
ment. This allowed relatively even coating of complex
surface topography. The gilded stub was taken from the
vacuum evaporator and transferred into the SEM speci-
men chamber. After pumping the electron column and
chamber down to 10° torr, study was undertaken.
Special equipment permits insertion of specimens up to
36 mm in size, but movements are greatly restricted and
there is no increase in the field of vision. A specimen 12 mm
square and 3 mm high can be handled with no restrictions
on adjustments.
MATERIALS
Figures 1 and 2 are of a Moorea, Society Island species
formerly known as “Charopa” modicella (Férussac,
Explanation of Plate 58
“Charopa” modicella (FERUSSAG, 1840)
Figures 1, 2: Faatoai Valley, Moorea, Society Islands. Bernice P Bishop Museum number 150377. Figure 1:
diagonal view onto
apex and early postnuclear whorls at 335 X magnification. Figure 2: enlargement of upper right area from negative of Figure-1 to dem-
onstrate presence of raster lines.
Ptychodon microundulata (SuTER, 1890)
Figures 3-6: Turanganui River, Haurangi Range, Wellington, New
Zealand. Dominion Museum, Wellington. Figure 3: General view
of shell at 110 X. Figure 4: Suture between nuclear and postnuc-
lear region at 1 100 X showing apical (upper left) and post-apical
(lower right) sculpture types. Surface debris on right side can be
used, to locate area photographed slightly above and to the left of
center in Figure 3. Figure 5: Sutural area at end of nuclear whorls,
1 100 X, showing tentative start of microradial sculpture and chan-
neled suture. Photograph taken at 18 whorl mark and showing
post-apical sculpture at the end of the first complete postnuclear
whorl. Figure 6: Detail of early apical sculpture at 3 350 X showing
irregularly folded periostracum between two radial ribs. Folds can
be matched with upper left portion of Figure 4 to establish size
change.
Plate 58
THE VELIGER, Vol. 12, No. 4 [Sotem] Plate 5
Figure 5 Figure 6
Vol. 12; No. 4
1840). It is an endodontid land snail belonging to the
subfamily Charopinae, but is classified in a currently un-
described genus. The specimen is from Faatoai Valley and
is Bernice P. Bishop Museum number 150377. Remaining
figures are of the New Zealand endodontid Ptychodon
microundulata (SuTER, 1890). Specimens were collected
by A.C. O’Connor in June 1949 from the Turanganui
River, Haurangi Range, Wellington and preserved as
dry specimens at the Dominion Museum, Wellington. I am
deeply indebted to the authorities of the Bishop Museum
and Dominion Museum for permission to study this
material.
RESULTS
Dissections of Pacific Island endodontid land snails had
revealed anatomically quite distinctive subfamilies whose
shell sculpture appeared to be virtually identical under
optical microscope examination. Initial use of the SEM
was in search of conchological features that would serve as
subfamily or generic identification criteria. Expansion of
this work into studies of functional aspects of shell sculp-
ture and investigation of endodontid shells from other
areas soon followed.
The Moorean specimen is tilted at about a 45° angle
from a horizontal view. Figure 1 shows the apex and part
of the first two postnuclear whorls in a shell measuring
3.25 mm in diameter with 44+ whorls. Actual width of the
first nuclear whorl, which is not completely shown in
Figure 1, is about 0.38 mm. Both apical and postnuclear
sculpture are typical for the Pacific Island Charopinae.
Ptychodon microundulata has a relatively complex and
highly photogenic surface sculpture (Figures 3 to 11). All
these photographs involve portions of a juvenile shell
measuring 1.38 mm in diameter with 33+ whorls. Photo-
graphs of the parietal lamella (Figures 12-15) were
taken from a slightly larger individual by breaking away
the palatal wall so that a direct view could be obtained of
the entire lamella. Optical examination at 100 had
shown that the juvenile showed essentially no worn spots
on the apex and postnuclear whorls. More information
could be obtained from this specimen than from larger
examples with obvious surface wear. Adult size for this
species is 1.71 - 1.97 mm, X is 1.86 mm, S. E. M. is 0.024
mm with 10 specimens measured.
Figures 4 - 6 and 8 - 10 were taken during one visit to
Alpha Research. Figures 3 and 7 were taken about two
months later. Despite the specimen having been kept in
a closed snap-lock plastic box, the concentration of par-
ticulate matter in Chicago air is obvious. More than half
of this debris accumulated after the initial study.
THE VELIGER
Payers 3)3)/|
Figure 3 at 110 indicates essentially what can be seen
under optical examination, although only a small part of
the shell would be in focus at any one time. Major shell
sculpture consists of prominent radial ribs on both the
apex (first 13 whorls) and postnuclear whorls. The radial
ribs are more widely spaced on the early apex, become
much more crowded on the last portion and then change
in appearance at the start of postnuclear growth. While
the presence of some kind of microseulpture between the
radial ribs can be seen, its frequency and nature cannot be
determined.
Although the texture of the interstices between the
apical radial ribs looks different from the texture between
the postnuclear radial ribs, no information is retrievable
at 110><. Under optical examination, data gathering
would stop. By switching the SEM to 1 100 and focus-
ing on a suture between the first apical whorl and first
postnuclear whorl (Figure 4), both microsculptures are
displayed. The postnuclear portion (right and lower half)
has a series of complex microradial riblets with enlarged
“beads” that are arranged in spiral rows. Apical sculpture
consists of irregular wrinkles and ridges with a “glassy” or
“greasy” surface appearance. At 3350 (Figure 6) this
apical sculpture is confirmed as consisting of a smooth
film (= periostracum) that is irregularly folded. Rota-
tion of the specimen to the point of demarcation between
apical and postnuclear whorls (Figure 5) and examination
at 1100 shows that tentative formation of the post-
nuclear micro-radial ridges begins a short distance before
the end of the periostracal film. Regular production of the
micro-radial ridges starts at the end of this folded film
and coincides with the increase in rib spacing noticeable
in the early postnuclear section of Figure 3.
Several other New Zealand endodontids and the Indo-
nesian-Pacific Island Discocharopa have the same type of
apical periostracum. An essentially equivalent type is seen
in several European taxa that were photographed in co-
operation with Dr. Gittenberger of Leiden. Interpretation
of this structure is somewhat difficult. In physical char-
acteristics, John A. Brown of Alpha Research informs me,
this is the same pattern seen in many dried paint or var-
nish films at equivalent magnifications. With this know-
ledge, a hypothesis of origin can be suggested. Both the
periostracum and calcified layer of the apical whorls are
formed during a late embryonic stage. This takes place in
liquid and the embryonic shell with firm calcification can
be removed from unhatched eggs. Thus solidification of
the calcium layer may occur while the periostracum is
still bathed in liquid. If hardening of the periostracal
layer was delayed until after hatching, then air drying of
the periostracal film could result in the irregular folding
Page 398
seen in the apical film. Differential frictional adherence
of the film at the start of drying would result in the devel-
opment of “tension folds” as it settled or shrank: into
position on top of the calcium layer. Possibly the initial
movement of the snail from the ruptured egg shell or
brushes against minute particles in the environment could
shift or distort the wet and pliable film, thus producing
the irregularities. Even rotational movements of the em-
bryo within the egg capsule might be enough to produce
the distortions. Determination of what is the time rela-
tionship between end of the apical sculpture and hatching
will require study of young that were preserved minutes
after hatching. While logic would suggest that hatching
would coincide with the start of postnuclear sculpture
formation, it is quite possible that it may precede hatching
for a few ridges during the period when the embryo is
actively moving but has not yet broken through the egg
shell. This could be determined very easily with the SEM
through study of pre-hatched, hatching, and recently
hatched shells.
Returning to the postnuclear sculpture, at 335 x (Figure
7) it is seen to consist of high major ribs that stop on
the suture edge. From 3 to 7 micro-radial riblets lie in
the trough between each pair of major ribs on the early
sections. They also are present on both the sides and tops
of the major ribs. Major ribs are 4 to + as wide as the
interval between each pair and the spacing varies slightly
from rib to rib. There are fewer micro-riblets between
crowded than between widely spaced ribs. This leads to
the conclusion that depositional timing of the micro-rib-
lets and major ribs is independent. That is, production of
a major rib is not started after “x” number of micro-rib-
lets, but that control of these two sculptures is separate.
Data on this aspect will be presented elsewhere. Optical
examination of the shell had permitted counting the
major ribs on the body whorl with an accuracy of +1.
Knowing that the shell diameter was 1.38mm, simple
multiplication by 7 produced the shell circumference' of
THE VELIGER
Vol. 12; No. 4
4.34 mm. Division of the 124 ribs on the body whorl
by the circumference in mm established there are an
average of 28.6 major ribs per mm on the body whorl
periphery.
By shifting to the body whorl periphery about 4 whorl
behind the aperture (Figures 8 to 11), it was possible to
examine structural details of the postnuclear microsculp-
ture. The basic orientation is looking down onto the body
whorl top and curvature of the periphery with a slight
angling towards the aperture. Whenever repetitive surface
sculpture is present, viewing around a curve permits simul-
taneous inspection of the sculpture elements from various
angles. This greatly aids interpretation and allows mental
stereoscopic reconstruction of complex topography. In
Figure 8 the magnification of 1 100 shows about 0.15
mm of body whorl periphery. Each major rib is seen to
have a single, high, posteriorly recurved, slightly sinuated
lamellar extension. Inspection at 3350 in Figure 9
shows that this structure arises from the top of a micro-
radial and that its surface shows traces of very irregular
grooving. As the snail moves through leaf litter, this
lamellar extension would brush against fine particles in
the environment. Minute grains of rock and soil would rub
against this reflected edge. As the snail moved forward,
sharp edges on the miniature boulders would score or
abrade the surface, producing the irregular grooves seen
on its outer edge. This lamellar extension is not present
during the first postnuclear whorl (Figures 3 to 5) and
only weakly developed on the second postnuclear whorl
(Figure 7). Most major ribs show partial evidence of
minor extensions on the microradials just before and just
after the major extension (Figures 8 to 10). An under-
' Although the actual measurement should have been one volution
of the logarithmic growth curve to establish periphery length, this
simple calculation is an adequate approximation. Rib counts are
the same at the suture and periphery of a whorl although the
linear distance is not the same. Bias is essentially the same for
every specimen and is not considered significant.
Explanation of Plate 59
Ptychodon microundulata (SuTER, 1890)
Figures 7-11: Turanganui River, Haurangi Range, Wellington,
New Zealand. Same specimen used in Figures 3 - 6. Figure 7: Early
apical and two whorls of post-apical sculpture at 335 %. Note suture
and “beaded” appearance of microradials except at curve into
suture. Figure 8: Periphery of body whorl, one-quarter whorl behind
aperture, at 1 100 X. Note change in appearance of microradials as
the curvature increases. Figure 9: Peripheral view between two
major ribs at 3 350 X showing change in microradial buttress form
and structure of microsculpture. Figure 10: Detail of microsculp-
ture on periphery at 11 100 X. Note change in buttress height and
shape on ascending side of major ridge at right margin of photo-
graph. Figure 11: Two “beads” on a microradial ridge at 33 500 X
showing the ascending buttress structure (apical direction to left
of figure, apertural direction to right). Figures 9 to 11 are enlarged
details from the area shown in Figure 8.
Tue VELIGER, Vol. 12, No. 4 [Ss ]
7 oLEM] Plate 59
Figure 11
Vol. 12; No. 4
THE VELIGER
Page 399
standing of this becomes possible after analysis of the
microsculpture.
At magnifications of 1 100 or less, particularly when
viewing at essentially a vertical angle (Figures 4, 5, 7)
the microelements appear as thin radial ridges with a
series of “beads” arranged in spiral rows. Part of this is
an artifact of electronic scanning. In Figure 8 at 1 100,
the lower left part of the photograph shows essentially a
vertical view. Note the beaded appearance of the sculp-
ture. Since the intensity of a scanned spot depends upon
both the angle at which the electron probe hits the sur-
face and the distance secondary electrons must travel to
the detector, a higher spot in direct path of the electron
probe will register much brighter than a lower surface
that is hit at an extreme angle by the probe. The elevated
“beads” thus register very brightly, while the sides and
significantly lower “trough” areas appear as dark spots.
As the angle of the electron probe increases towards the
center of Figure 8, the complexity of this sculpture is
revealed. At the higher magnifications of 3 350 (Figure
9), 11 100 (Figure 10), and 33 500 (Figure 11) this
can be interpreted. A network of spiral and radial ele-
ments comprise the microsculpture. The elevated beads
are connected radially (bottom to top in Figure 10) by a
narrower (Figure 11) buttress with rounded upper mar-
gin that gradually rises up on one side of the elevated
point and descends on the other. As these radial rows
approach the upsurge on the apical side of a major ridge
(right part of Figure 10), the radial buttress narrows and
becomes elevated above the rounded beads. Finally the
uppermost row on the major rib surface extends into the
reflected lamellar extension.
Spiral elements (left to right) in this sculpture consist
of a low ridge that sweeps up on the apical side in an
accelerating curve to meet the radial buttress and extends
above it as a “bead”. Generally it drops off extremely
rapidly on the apertural side of the bead, but Figures 9
and 11 show that a more gradual drop is possible. At the
bottom of the trough between each pair of spiral butt-
resses are a few irregular folds. Figure 11 shows that these
are sometimes present on the elevated buttresses. A pos-
sible explanation is that these are periostracal tension
folds. Growth proceeds from left side to right side in these
photographs, so that the less elevated portion of the
microradial sculpture is deposited first. With subsequent
extreme elevation of the radial buttress, even slight de-
parture from the exact template relationship of the peri-
ostracum and underlying calcium deposition would create
stress in the periostracal layer. Hence the folds in this
region could be analogous to the apical folds, although at
a much lower level of visibility.
Obviously the use of 300 to 30000 SEM magnifica-
tions permits gathering infinitely more data concerning
the composition and growth patterns of shell sculpture
than is available using optical instruments. The previous
standard of description might refer to such a species as
“radiately costulate” or “major growth riblets about 21
per mm on body whorl with a few finer riblets between”.
Now it is possible not only to study the exact physical
structure of such sculpture, but to pose questions con-
cerning its mode of formation, to measure its relative
prominence, and to compare functioning effectiveness of
this sculpture in different taxa.
Equally important data can be obtained concerning the
structure of apertural barriers in small land snails. These
occur in many different land snail families, sometimes in
adults and sometimes in juveniles, but are particularly
common in endodontid land snails. Figures 12 to 15 illus-
trate details on the posterior half of the major parietal
lamella in Ptychodon microundulata. Previous illustrations
(Surer, 1890, plt. XIV, fig. 4c; Surer, 1915, plt. 27, fig.
8c; and Ciro, 1969, p. 197, fig. 12) show very little
detail, either portraying the lamella as smooth or with
serrated edges. There are 3 rows of large denticles on the
expanded margins of the lamella, with the denticle points
directed towards the aperture (Figure 12). Two rows
are on the lower margin, one on the upper. Even at 335
(Figure 12), but particularly at higher magnifications
(Figures 13 to 15), one can see that a thin transparent
film connects the points. This is dried mucus. Frequently
such mucous sheets are found on the shell surface or partly
covering apertural denticles in specimens that were col-
lected alive, but then simply air dried and kept in a collec-
tion tray. Under these circumstances, the mucous traces
caused by snails crawling over each other in the collecting
vial, or the patch of mucus left as the snail retreats into
the shell before dying of desiccation, hardens into a sheet
that is almost impossible to loosen by soaking or ultrasonic
cleaning. Specimens that were drowned and preserved in
alcohol have loose mucous coatings that are readily
cleaned. Since this shell was a live collected, air dried spe-
cimen, the mucous sheet was present and could not be
removed.
Irregularity in angulation, size and position of the den-
ticles is evident. At 1100, study of the lower two rows
(Figure 13) shows that the interstitial area contains mi-
Page 400
nute crystalline structures. This is particularly clear in the
lower left portion. In the central area of this photograph,
the overlapping mucous sheet obscures details. Shifting
study to a lateral view of the upper denticle row in Figure
14 at 1100 and Figure 15 at 3350 indicates the na-
ture of the growth pattern. The denticles are simply
elongated prismatic aragonite rods. Figure 15 shows that
the low crystalline structures in Figure 13 are shorter
crystals that parallel the growth of major denticles. In the
upper left part of Figure 15 the surface texture is the same
as prismatic growth area surfaces figured by WISE & Hay
(1968b, p. 422, fig. 8). This is the smooth interdenticle
row area shown in Figure 12. Note the slight discontinuity
between this prismatic growth surface section and the
aragonite rods. This is typical of such an interface and has
no special significance. Other lamellae on the columellar
and palatal walls differ in having smooth surfaces, even
at 3000 magnification.
SUMMARY
Scanning electron microscopy permits whole specimen
photography of mollusks up to 7 mm in size, with the
image filling a 3534 inch negative. Since the depth
of field obtainable is several times greater than the height
or width of the picture, all areas will be in sharp focus.
Parfocal changing of magnification between 20 and
50 000>, combined with the ability to tilt, shift and rotate
the specimen being observed, permit examination of suc-
cessively finer detail, stereoscopic photograph preparation
and examination of the same structure from different
angles.
As a result, questions that previously could not be asked,
now may be answered quickly. Molluscan shells present a
frozen record of growth during the life of that individual.
Details of this past are readily visible by electronic anal-
ysis. Ptychodon microundulata is presented as a partic-
ularly fine example of complex sculpture, together with
some of the questions and hypotheses that result from
examining this species.
THE VELIGER
Vol. 12; No. 4
LITERATURE CITED
BLAKER, ALFRED A.
1961. Basic lighting for shell photography.
3 (3): 69-72; plt. 12; 3 text figs.
Cumo, Frank M.
1969. Classification of the New Zealand Arionacea (Mollusca:
Pulmonata). II. A revision of Charopa subgenus Ptychodon.
AncEy, 1888. Records Domin. Mus. 6 (14): 175 - 258;
11 plts.; 34 text figs. (23 May 1969)
Ersen, H.K., G. Fiays « A. SIEHL
1968. Uber die Schalenstruktur von Monoplacophoren.
Akad. Wiss. Lit. Abhandl. Math.-Naturwiss. K]., Mainz 1968
(1): 1-24; plts. 1-17
Hay, WiLiiAM W. & Puiuie A. SANDBERG
1967. The scanning electron microscope, a major break-through
for micropaleontology. Micropal. 13 (4): 407 - 418; plts.
The Veliger
(1 January 1961)
il, 2 (October 1967)
SoLem, ALAN
1969. ... in his dim, uncertain sight. Bull. Field Mus.
Nat. Hist. 40 (3): 7-9; 12 figs.
SuTER, HENRY
1890. Descriptions of new species of New Zealand land and
fresh-water shells. Trans. Proc. New Zealand Inst. for 1889,
22: 221 - 230; pits. 14, 15 (May, 1890)
1915. Manual of the New Zealand Mollusca. Atlas of
Plates, 72 plts. Wellington (John Mackay) .
Taytor, Joun D., W. J. Kennepy « A. Hatu
1969. The shell structure and mineralogy of the Bivalvia. In-
troduction. Nuculacea- Trigonacea. _ Bull. British Mus. (Nat.
Hist.), Zool. Suppl. 3: 1-125; 77 figs.; 29 plts.
THompson, Tuomas E. « H. E. Hinton
1968. Stereoscan electron microscope observations on opistho-
branch radulae and shell-sculpture. Bijdr. Dierk. 38: 91
to 92; plts. 1-4
WIsE, SHERWOOD W.
1969. Study of molluscan shell ultrastructures. In: Scanning
Electron Microscopy/1969, Proc. 2.4 Ann. Scanning Electr.
Microsc. Symposium, pp. 205-216; 23 figs. Chicago, Illinois
(IIT Research Inst.) (April 1969)
Wist, SHERWoop W. & WILLIAM W. Hay
1968a. Scanning electron microscopy of molluscan shell ultra-
structures. I. Techniques for polished and etched sections.
Trans. Amer. Micros. Soc. 87 (4): 411-418; 8 figs.
1968b. Scanning electron microscopy of molluscan shell ultra-
structures. II. Observations of growth surfaces. Trans. Amer.
Micros. Soc. 87 (4) : 419 - 430; 17 figs.
(1 April 1969)
Explanation of Plate 60
Ptychodon microundulata (SutTER, 1890)
Figures 12-15: Parietal lamella from adult specimen in same set
as juvenile photographed in Figures 3 to 11. Figure 12: Posterior
portion of parietal at 335 X, viewed slightly from below. Figure 13:
Double rows on lower edge of lamella at 1 1000 X. Note fine crys-
talline rods visible between major denticles. Figure 14: Lateral view
of upper row at 1 100 X. Figure 15: Lateral view of upper row
at 3 350 X. Note mucous film and aragonite rods.
TuHeE VELIcER, Vol. 12, No. 4 [Sorem] Plate 60
Figure 14 Figure 15
eh
if
ie
Vol. 12; No. 4
THE VELIGER
Page 401
The Effect of Wave Impact
on Some Aspects of the Biology of Sea Mussels
J. R. E. HARGER
- Department of Biological Sciences, University of California at Santa Barbara '
(9 Text figures; 15 Tables)
INTRODUCTION
THE ABSENCE OF a simple wave impact measuring device
has hampered the efforts of ecologists in comparing con-
ditions among different intertidal habitats for some time.
Moore (1935) has made use of the percentage of the
year’s winds blowing into the angular aperture of a
locality; SourHwarp (1958) used the occurrence of
waves washing over the breakwater at Plymouth, coupled
with known wind records, to generate a wave index;
BALLANTINE (1961) has outlined a biologically defined
exposure scale for the comparative description of rocky
shores; and finally Erion Jones & DOoMETROPOULOS
(1965) have used an apparatus consisting of a drogue at-
tached to a spring dynamometer to record drag produced
by passing waves.
(I) MEASUREMENT or WAVE IMPACT
In investigating the effects of wave action on the biology
of two species of mussels (Mytilus edulis Linnagus, 1758
and M. californianus Conrap, 1837) on the coast of
Southern California, I made use of a device for measur-
ing wave impact. This consisted of a smooth 6 inch long
nail, a spring steel “c” clip, and a metal plate of 1/16
inch gauge steel. A 4+ inch diameter hole was drilled in
the center of the plate which was then passed up the
nail shaft until it rested against the head. A “‘c” clip was
then threaded up the shaft of the nail to the bottom of
the plate so preventing the plate, when released, from
falling down the nail (Figure 1). 'The nail was then fixed
into an “intertidal” rock or pier piling so that the head,
and metal plate supported by the “c”clip, projected into
' Present address: Department of Zoology, University of British
Columbia, Vancouver 8, British Columbia.
the prevailing wave motion. Wave action, during one
tidal period, then forced the metal plate and “‘c” clip
down the shaft of the nail. The distance through which
the plate is moved depends on 3 factors: size of the plate,
frictional resistance afforded by the “‘c” clip, and strength
and frequency of wave impact.
The first 2 factors can be controlled; all that is
necessary is to insure that such a combination of plate
size and clip strength is used, at a particular geographic
location, that the plate will not be forced completely down
the nail during one tidal period. The force exerted by
heavy storms can be estimated with this device, simply
by making the plate very small.
metal — Z?
plate
6” nail
Figure 1
Wave impact measuring device
Page 402
The force of wave impact for one (or more) tidal
cycles is calculated in the following manner:
a = area of the plate in cm’
f— force required to overcome the friction of the
“c” clip, measured by pulling it down the nail with
a spring balance (in kg)
d= distance the plate is moved (in cm)
Wave force = (f/a)d. kg-cm/cm?
If the wave force is to be compared at several locations,
nails must be set out at each place within the same tidal
period. (I found that an individual “c” clip could be
used only 2 or 3 times and that the nail shafts must be
kept free of burrs and nicks.) Table 1 lists values ob-
tained for wave impact, over three 24-hour periods, from
5 locations.
These areas are all situated within a few miles of the
City of Santa Barbara, on the coast of Southern Califor-
nia (Figure 2). Stearns Wharf is located at the entrance
‘San Francisco
se UCSB Santa Barbara
Figure 2
Location map showing Santa Barbara and study areas
1. Ellwood Rocks 2. Ellwood Pier 3. Goleta Point
4. Stearns Wharf 5. Carpinteria
of the Santa Barbara harbor and is quite sheltered. Ell-
wood Pier (property of Signal Oil and Gas Co.) is a
large pier, almost $ mile in length, situated some 14
miles west of Santa Barbara, on open sandy shore. The
steel pilings of this pier extend from the intertidal surf
zone out to a depth of about 40 feet. These pilings sup-
port large intertidal clumps of mussels which are com-
prised of both Mytilus californianus and M. edulis. Ell-
wood Rocks are part of the shore line immediately
adjacent to Ellwood Pier, and Goleta Point is an out-
cropping of rock just below the campus of the University
THE VELIGER
Vol. 12; No. 4
of California at Santa Barbara (10 miles west of the City
of Santa Barbara). These points were chosen for easy
access and because a series of exposures ranging from
sheltered (Stearns Wharf) to exposed (Goleta Point)
were obtained. oa
During the recording period, winds were light (5-8
knots on-shore) but a heavy swell was running producing
breakers between 4 and 5 feet in height on the local sand
beaches. From Table 1, it appears that the force of wave
impact suffered by Ellwood shore is 4-5 times that in-
curred by the outer end of the adjacent Ellwood Pier.
Within the pier itself, a particular sheltered position (in-
take pipe) received less than 4 as much force as nearby
open pilings and Goleta Point experienced some 13 to
18 times more wave impact than the Ellwood shore.
The recorders at Ellwood Shore were driven into the
front face of a 3 foot high hemispherical boulder (8 feet
in diameter) which projects from the relatively gently
sloping lower mid-littoral (the recorders in all locations
were placed in the lower mid-littoral). The recording site
at Goleta Point was a 5 foot high vertical rock face situ-
ated in the lower mid-littoral region. This rock face is so
placed that a swell hitting it at half tide may throw spray
to a height of 12 to 15 feet.
These measurements indicate that the impact force ex-
perienced by any group of organisms on an exposed shore
line depends on the strength of the waves, on the aspect of
the rock face to which they are attached, and on whether
there is a beach, or other rocks, to seaward which can
absorb some of the force before the waves reach the or-
ganisms. There is a large difference between the exposure
value obtained from Stearns Wharf (sheltered) and that
from Ellwood Shore (exposed) (see Table 1). However,
on the open shore, rocks in different positions (Ellwood
Rocks and Goleta Point), are exposed to wave impact
forces which differ from each other by as much as the
difference between the sheltered harbor (Stearns Wharf)
and the stretch of exposed coast (Ellwood Rocks) impact
values. Taking into account the difference between the
front (seaward) and rear (landward) faces of intertidal
rocks, it is clear that the intensity of wave impact on an
exposed intertidal shore must vary greatly. Wharf pilings
projecting from natural intertidal regions suffer more frora
wave impact than deep water pilings. This difference
between deep and shallow water pilings has also been
reported by Bascom (1964). On pilings, in general, wave
impact is greatest at the mid-tidal level and less at higher
and lower levels.
A method similar to that reported here of recording
the pressure exerted by waves against structures was used
by Thomas Stevenson in 1842. Bascom (op. cit.) reports
Vol. 12; No. 4
THE VELIGER
Page 403
Table 1
Mean wave impact values obtained on three days from
four different sites
(The unit of measurement is kg—cm/cm?)
12 Dec. 1966 13 Dec. 1966 15 Dec. 1966
Mean Mean Mean
Wave Wave Wave
Place Impact Sample Impact Sample Impact Sample
Values Size Values Size Values Size
Stearns Wharf, Santa Barbara Harbor 0.012 3 0.014 3 0.022 3
Ellwood Pier (Intake Pipe) 0.048 4 0.055 1 0.060 1
Ellwood Pier (Outer Pilings) 0.105 2 0.152 5 0.199 4
Goleta Point <3.769 6 9.160 3 9.187 3
Ellwood Rocks 0.495 5 0.590 5 0.766 6
that the instrument used “... consisted of a plate six
inches in diameter facing into the waves, mounted on a
stiff horizontal spring. Behind the spring was a rod held
by a friction grip in such a fashion that it would move as
the plate moved but remain at the maximum distance
which the plate pushed it. As each increasingly large
wave impacted against the plate, the rod would be pushed
to a new position. The distance moved times the spring
constant gave the maximum wave (single) force exerted
on the plate ...” The modern “professional” pressure
gauge which is now used by engineers to measure the
force exerted by waves on pilings, piers and shoreline
structures is described by Bascom (op. cit.) as “consisting
of a stack of thin plates of tourmaline crystal set in a strong
metal case. When subject to pressure, this gage produces a
small charge of electricity which can be amplified ...”
In the same book, data which indicate that there is a
substantial increase in the force exerted on a piling as
waves change from swell into breakers and then foam-
lines are presented. Bascom also comments that “measure-
ment of wave forces on pilings is complicated by the con-
tinual reversal of direction of the water as the crest
moves in one direction and the trough moves in the
UNS, goo
Table 2
Comparison of growth increment data:
Mytilus californianus and Mytilus edulis from exposed
and sheltered cages set at the mid-tide level on the outer
end of Ellwood Pier. The mussels used in the experiment
were all originally between 3 and 4cm long. Growth
period was from December 1966 to March 1967
All measurements are in centimeters
Note: one asterisk (*) indicates significance at the 5% level, two
asterisks (**) significance at the 1% level and three asterisks (***)
significance at the 0.1% level. Abbreviations of statistical terms are
those used by J. C. R. Li (1964).
§ E S 2
2 form.)
© 12} EN F
a
Group a as A 3,155
1. Mytilus edulis from plastic cage 0.94 0.57 33 49.01%
2. Mytilus edulis from wire cage 0.34 0.27 44
3. Mytilus californianus from plastic cage 1.53 0.72 42
4. Mytilus californianus from wire cage 1.44 0.56 40
F
1,155
Individual D. FE Tests
1.2/3.4 108.56***
3/4 0.55N.S.
1/2 20 Dee
Page 404
THE VELIGER
Vol. 12; No. 4
The advantages of the wave impact measuring device
described in this paper lie chiefly in its simplicity and low
cost. By using several instruments at several locations an
ecologist can get a quick (although perhaps somewhat
rough) estimate of the wave impact force acting on dif-
ferent parts of the intertidal region.
(II) EFFECTS or WAVE IMPACT
on THE BIOLOGY oF
Mytilus edulis ann Mytilus californianus
(A) Growth
The effect of wave impact was investigated by placing
equal numbers of each species of mussel (individuals 3 - 4
cm in length) in open mesh wire cages (exposed treat-
ment) and in cages of partly occluded sheet plastic (pro-
tected treatment) and recording subsequent growth. The
first type of cage was constructed from 4 inch mesh (1.27
cm) galvanized hardware cloth and was cylindrical in
shape (diameter 7 inches {17.78 cm}) with a height of
8.5 inches (21.5 cm). The various components (wire sec-
tions, etc) used in construction were laced together with
braided nylon cord and the entire structure then coated
with epoxy resin. This treatment served to give rigidity
to the nylon binding and at the same time to cut down
any leaching of zinc ions which might affect the enclosed
mussels. The second cage was made of plastic kitchen
colanders (10 inches {25.4cm}) in diameter placed
face to face and lashed together at the edges (the maxi-
mum diameter of the holes in a colander was } inch
{0.963 cm}).
The cages were suspended at the mid-tide level on the
extreme outer edge of Ellwood Pier. After 4 months (De-
cember, 1966 to March, 1967), the growth increment
Table 3
Comparison of growth increment data:
Submerged Mytilus edulis from a plastic cage, a wire
cage and a naturally occurring clump of mussels growing
on an electrode cable
Original shell length has been used as the independent variable
and growth increment as the dependent variable. All measurements
were made in centimeters.
2 Fh BSS 2
ag a, 2 th ASS
Group on Regression Equation z a} = 3 ti OS
7) aX
1. Mussels from 27 Y=0.005+0.15X 3.04N.S. 1.011 1.96N.S.
clump on cable
2. Mussels from 33 Y =0.045+0.17X 1.153
wire cage
3. Mussels from 42 Y=0.491 + 0.08X 1.046
plastic cage
Table 4
Comparison of growth increment data:
Submerged Mytilus californianus from a plastic cage, a
wire cage and from a naturally occurring clump of mussels
growing on an electrode cable
Original shell length has been used as the independent variable
and growth increment as the dependent variable. All measurements
were made in centimeters.
ay oat B hgh: rege Mae
Group a & Regression Equation i) ies 2 g i ae
<2x
1. Mussels from 11 Y =0.74+0.023K 0.37N.S. 0.879 1.08N.S.
clump on cable
2. Mussels from 28 Y =0.64+0.072X 1.092
wire cage
3. Mussels from 43 Y=0.52+0.089X 1.072
plastic cage
a —————0———_—_—_—_—_—_—_—_—
Vol. 12; No. 4
THE VELIGER
Page 405
for each mussel was recorded. It was found that the
Mytilus californianus populations grew much faster than
those of M. edulis (Table 2). There was no significant
difference in the growth rate of M. californianus from
the different cages, but M. edulis from the wire cages
grew 4 the amount shown by the animals in the plastic
colander. This difference was attributed to the effect of
wave impact which presumably acted more severely on
the mussels in the wire cage (the openings in the wire
mesh account for approximately 92% of the cage surface)
whereas the walls of the colanders, because they were
constructed of sheet plastic containing relatively small
holes (the holes account for approximately 15% of the
cage surface) would diminish the impact of breaking
waves. The fact that no significant difference in growth
can be detected between M. californianus from the shel-
tered and the open cages would seem to indicate that this
species is not sensitive to the exposure differences between
the two cages. A further test was conducted at the same
time to determine the effect these two cages might have
on the growth of both species of mussels when placed
below low water where the effects of wave impact are
absent. An additional comparison was made in this case
between the two treatments using caged mussels and
mussels growing naturally within submerged clumps at-
tached to insulated cables suspended from the pier.
Growth for each species was not detectably different for
any of these treatments (Tables 3, 4), which indicates
that caging itself does not hinder growth under these
conditions.
(B) Size
Figure 3 illustrates the relationship between the maxi-
mum size of Mytilus edulis occurring naturally at several
localities and the corresponding wave impact values. It
can be seen that small mussels are associated with high,
and large mussels with low values. This relationship is
consistent with what one might predict from knowledge
of the force required to remove different sized individuals
of M. edulis from a rock surface. This force was recorded
by fastening either a battery clip to, or looping a piece
of cord around a mussel and then hooking a spring scale
through the clip or a loop on the cord and noting the
force required to dislodge the mussel as the scales were
pulled firmly away from the rock. Analysis reveals (Fig-
ure 4, Table 5) that the force necessary to tear loose the
two species of mussels was significantly different, with
M. californianus requiring more force when similar sized
mussels are compared. Data relating to a third species
of mussel, Septifer bifurcatus (Conrap, 1837) are also
included with the analysis given in Table 5 and illustrated
in Figure 4. These mussels occur intertidally on the Santa
Barbara coast but are physically quite small — the maxi-
mum length attained being about 3.5 - 4.5m long. The
(cm)
Maximum Size of Mytilus edulis
0.0 0.10 1.00
10.00
Wave Force (kg — cm/cm?)
Figure 3
Relationship between wave impact values and the maximum size of
Mytilus edulis occurring at the measurement sites
1. Stearns Wharf 2. water intake pipe at Ellwood Pier
3. outer end of Ellwood Pier 4. Ellwood Shore 5. Goleta Point
force required to remove an individual from a rock face
is greater than that required to remove a similarly sized
M. californianus individual. (Populations of Septifer are
found intertidally only in exposed conditions. )
The relationship indicated for Mytilus californianus
(Figure 4) is strictly linear, even for the largest mussel
tested (10 cm long), but the curve obtained for M. edulis
flattens out once a length of 4-5 cm is reached, so that
the force required to remove a 10 cm individuai is the
same as that required to remove a 5cm mussel. The
larger mussels used to provide points on this curve all
came from various positions on Ellwood Pier, some from
pilings and some from large subtidal mussel clumps which
had formed on zinc electrode cables hung from the pier.
Since these cables do not touch the bottom, starfish are
unable to find and attack the mussel clumps, so, in
Page 406
26.0
Force (kg)
Shell area (cm?)
Figure 4
The force to remove mussels from the open shore plotted against
a correlate of shell area (length X breadth).
[Shell measurements in centimeters]
Septifer bifurcatus represented by solid triangles, Mytilus californi-
anus by solid circles and Mytilus edulis by dashes. Crosses indicate
values for both M. californianus and M. edulis. Regression lines are
fitted by least squares (see Table 5)
the absence of predation these mussels may reach an
extremely large size in the relatively quiet water (M.
californianus up to 30cm and M. edulis up to 15cm
long). Only mussels unencumbered by the byssal threads
of others were used in the tests; even so, it might be
argued that since the large mussels came from a complete-
ly different environment from that of the shore mussels
then the relationship in Figure 6 is not representative of
a shore population and that it is therefore invalid to
draw inferences from it reflecting the disposition of such
populations. It was however impossible to find large in-
dividuals of M. edulis on the shore, therefore, sub-tidal
specimens were used. In an attempt to counteract the
THE VELIGER
Vol. 12; No. 4
above objection a laboratory test was performed, the
results of which are presented in Table 6 and Figure 5.
Individuals of both species were placed for 3 weeks on
the bottom of tanks containing running sea water. The
force required to remove them at the end of this time was
then recorded. The relationship is essentially the same
_as that obtained by using the naturally set animals, how-
ever, since these mussels established fewer byssal threads
the dislodging force was generally lower.
Since small Mytilus edulis survive wave impact with
apparent success, a comparison of the two species involv-
ing animals of up to 4.cm in length was also made, The
results show clearly that even at this size M. californianus
holds on more strongly than does M. edulis (Table 7).
From these results one might predict that only small
individuals of Mytilus edulis could survive in “exposed”
situations and that conversely large specimens (5 cm and
over) would be likely to occur in sheltered places such as
harbors, etc. This is borne out by observations along the
coast of Southern California. Mytilus californianus on
the other hand is able to attach itself such that a
relatively constant force (in proportion to its shell area)
Shell Area (cm?)
Figure 5
Force to remove mussels from laboratory tanks (constructed from
resined fibre glass) plotted against a shell area correlate (length <
breadth). @ — Mytilus californianus; O-—M. edulis.
Shell measurements in centimeters
Vol. 12; No. 4 THE VELIGER Page 407
Table 5
Comparison of the force required to remove different
sized, naturally set, individuals of Mytilus californianus,
Septifer bifurcatus and Mytilus edulis
A measurement correlated with shell area (length X breadth) has the shell valves (from the dorsal to the ventral edges) : length was
been used as the independent variable and force required to remove measured as the maximum distance between the anterior and pos-
mussels (measured in kilograms) the dependent variable. terior edges of the valves, i. e., from hinge(umbo) to siphon regions.
The breadth measurement was made across the broadest part of All measurements were recorded in centimeters.
Bao
4 OG sy ee x
e ee uae eer | Oa! Terk
rou 3 egression Equation wWeWN oh av
2) w g q n z S 4
1. Septifer bifurcatus 32 Y=1.71+1.14X% 134.28*** 14.513 138.01***
2. Mytilus californianus 111 Y=2.14+0.58X 8.768
3. Mytilus edulis 125 Y=3.00+0.10X 4.171
F
Individual D. F Test on B 1,262
1/2 2.50N.S.
2/3 262.46***
Individual D. F. Tests on Adjusted Means
1/2 4.03*
2/3 267.95***
Table 6
Comparisons between the forces required to remove A correlate of shell area (length breadth) has been used as the
different sized individuals of Mytilus edulis and Mytilus independent variable and force required to remove the mussels
B . Frontech : ll Alsip En (measured in kilograms) as the dependent variable. Length meas-
californianus a ter pt SPECIES WERE. OWE ESO aUae urements made on the mussels were in centimeters.
the inside of laboratory tanks
Pid
a) n 3 S a
£8 Bet BEX wh
Group Fie Regression Equation = a Saat | =
2 <Z SA
1. Mytilus edulis 63 Y=0.40+0.024X 1A a12 ene O60) a3 2.08%e%
2. Mytilus californianus 16 Y =0.20+0.090X 1.529
Table 7
Comparisons between the forces required to remove A correlate of shell area (length > breadth) has been used as the
independent variable and force required to remove the mussels
individuals of Mytilus edulis and Mytilus californianus ee :
Senn dhe ouan drone when bath are less dhan 997 Gan (measured in kilograms) the dependent variable. Length measure-
P ; : ments made on the mussels were in centimeters.
in length
Means at
Slopes
F
1,58
Adjusted
&
a
Group Fie Regression Equation
1. Mytilus edulis 20 Y =0.26 + 0.604X 3.75N.S. 2.064 HOO Re
2. Mytilus californianus 42 Y =0.28 + 1.183% 3.253
Page 408
THE VELIGER
Vol. 12; No. 4
is necessary to detach it, consequently all sizes (at least
those up to 10cm long and probably longer) are able
to withstand wave impact equally well and thus no
size limitation seems to be imposed on this species by
wave action.
(C) Body Weight
Body weight is a further characteristic of mussels which
varies with the force of wave impact. This was demon-
strated by collecting a group of mussels from each area in
which wave impact data were recorded. After recording
individual dry body weights, an adjusted (for shell length)
mean dry body weight for each group was determined.
Figure 6 shows that the body weight of Mytilus edulis
varies inversely with the amount of wave impact to which
it is exposed.
0.8
0.7
0.6
0.5
Adjusted Mean Dry Body Weight (gm)
5 Io 20
Wave Force (kg — cm/cm?X 10?)
3o40
Figure 6
Plot of the square root of dry body weight of Mytilus edulis
(adjusted to a shell length of 4.037 cm) against wave force values.
The latter values were obtained from four geographical positions
(Positions 1 to 4 in Figure 3). Values obtained on three different
days (vy, mw, @ ) are plotted for each position. The bar repre-
senting variation of body weight is proportional to twice the
standard error of the mean on each side
It was not possible to obtain such a precise relation-
ship for Mytilus californianus — it seems that in spite of
the amount of wave impact experienced, this species
maintains a relatively constant body weight. This is not
to say that the body weight of M. californianus from
different areas is constant, because this is not so. A large
amount of variation exists here and the closest thing to
a general statement which can be made with regard to
this (based on numerous separate samplings from a num-
60 80
ber of areas) is that the body weight of M. californianus
from an exposed area (Ellwood Shores) is likely to be less
than that from a relatively sheltered area (Ellwood Pier
pilings) (Table 8), but this is not always the case.
(D) Production of Check Rings
Both species develop “check rings” the number of
which is directly related to the length of the shell. This
may be a structural modification conferring additional
strength to the shells. In Mytilus californianus these rings
are numerous and prominent throughout the length of
the shell, and give it an appearance much like finely
corrugated iron. The shells of M. edulis on the other hand,
usually have few rings and appear quite smooth in com-
parison to those of M. californianus. For M. californianus
the number of check-rings per unit length of shell varies
51.0
44.0
37.0
isc)
S
°
Number of Check Rings
e)
16.0
0.00 0.80 1.60 2.40 3.20 4.00 4.80 5.60
Shell Length (cm)
Figure 7
Plot of the number of check rings against shell length for Mytilus
californianus taken from a clump of mussels at Carpinteria reef.
Solid squares represent mussels from the inside of the clump and
hollow squares, mussels from outside the clump. Regression lines
are fitted by least squares (see Table 9)
Vol. 12; No. 4
THE VELIGER
Page 409
Table 8
Comparison of dry body weight data:
Mytilus californianus from two “exposed” positions,
Ellwood Shore and Carpinteria Reef and from two
“sheltered” positions, Stearns Wharf and Ellwood Pier
The independent variable is shell length (measured in centimeters)
and the dependent variable is body weight (in grams). All body
weight measurements have been transformed by taking the square
root of the variable.
#19
a o ih & g < = S
EN ; GaN BR mA
Group 3 Regression Equation a Ge = S N oa)
1. Ellwood Shore 67 Y=-0.17+0.179X 6.34* 0.816 40.75***
. 2. Carpinteria Reef 74 Y =-0.27 + 0.233% 1.008
3. Ellwood Pier 37) Y =-0.21 + 0.238X 1.097
4. Stearns Wharf 30 Y =-0.26 + 0.226% 0.979
F
Individual D. FE. Test on B 1,220
1.2/3.4 6.37*
Individual D. F Tests on Adjusted Means
1.2/3.4 51.94%**
Table 9
Check ring comparison:
Mytilus californianus from inside and outside a clump
at Carpinteria Reef
The independent variable is shell length (centimeters) and the
dependent variable is the number of check rings.
Group
Sample
Size
1. Outside 64
2. Inside 60
Regression Equation
Y =-2.05+ 7.05%
Y= 0.87+9.00X%
no =)
4 © Gas o
Sen a
a — o =
2 Zsx
22.24*** 19.087 290.79***
27.884
Table 10
Comparison of the growth increase shown by two groups
of mussels growing inside a plastic colander placed below
low water at Ellwood Pier
Members of the first group were originally growing inside a clump
of mussels at Carpinteria Reef, those of the second group were
outside the same clump. The recorded independent variable was
growth increment (in centimeters).
Zs ©
S Ries! a
a P= for, v (=)
Group S a = Ba oer
nQ Z
1. Mussels from 0.59 0.33 38 0.05N.S.
inside clump
2. Mussels from 0.57 0.29 44
outside clump
with geographical location and even from different situ-
ations within a clump of mussels. It seemed that this
variation in check-ring frequency was in some way related
to the degree of disturbance, harassment or interference
that a particular animal had experienced during its
growth period.
The first step in testing this hypothesis was to obtain
mussels from inside and outside a clump from a moder-
ately exposed shore at Carpinteria Reef (a large flat
out-cropping of rock 10 miles east of Santa Barbara).
The analysis in Table 9 and Figure 7 shows that mussels
on the inside of this clump had more rings per unit length
than those on the outside. Mussels from the two samples
were then placed in separate plastic colanders which
were then lashed together face to face with a plastic
divider separating the two resulting compartments. This
Page 410
container was then suspended 3 feet below extreme low
water at Ellwood Pier.
After 4 months growth in this sheltered situation the
two groups were again examined. The analyses given in
Tables 10, 11, and 12 show that:
a) Growth rates of the two groups did not differ
from each other.
b) An apparent “basal” frequency of check ring
formation was established (i. e., rings within new
growth) which was the same for both groups (but
differed from the previous frequency established at
Carpinteria Reef).
c) This new frequency of ring formation (sheltered
position) was much lower than that of either of
the original groups (exposed position).
A further experiment at Ellwood Pier designed to com-
pare check ring frequency throughout the intertidal range
was established by shaving down a series of large mussel
clumps growing on the pier pilings until a single layer of
Table
Check ring comparison between two groups of mussels
growing inside a plastic colander placed below low water
at Ellwood Pier
THE VELIGER
Vol. 12; No. 4
mussels remained attached to each piling. This layer ex-
tended from the top to the bottom of the removed
clumps (approximately 6 feet or 1.9m). The mussels
comprising this layer were then allowed to grow for 5
months (February to July, 1966). A comparison was then
made between the frequency of rings laid down by the
mussels when they were confined to the insides of these
clumps (before I had removed the outer mussels) and
those laid down after they had been exposed. Results
indicate that, during the period of exposure, rings devel-
oped at a lower frequency per unit length than previous-
ly. Mussels higher up the pilings showed higher frequen-
cies than those lower down (both before and after being
exposed) (Tables 13, 14, 15). Since mussels from the
highest intertidal positions have the lowest growth rates
(Harcer, 1967) it seems reasonable to assume that they
are developing in positions in which “disturbance” fac-
tors (exposure, solar radiation, etc) act with greater
intensity and frequency than they would on subtidal
11
Members of the first group were originally growing on the outside
of a clump of mussels at Carpinteria Reef, while those of the second
group were inside that same clump. The independent variable is
growth increment (in centimeters) and the dependent variable is
number of check rings within the new growth.
8 A oR
Sur 38 i] ae
Group Regression Equation wa sh 3 A a
1. Mussels from 44 Y=1.72+3.53X 0.86N.S 3.485 3.63N.S.
outside clump
2. Mussels from 38 Y= 1.264 3.92X 3.222
inside clump
Table 12
Comparison of the frequency of check rings laid down by
Mytilus californianus when growing outside a clump of
mussels at Carpinteria Reef with that frequency shown
by the same mussels growing within a colander suspended
below low water at Ellwood Pier
The independent variable is shell length of the first group and
length of new growth for the second (in centimeters). The dependent
variable is number of check rings.
ae
Group Eu
7)
1 Mussels outside 64
clump Carpinteria
. 2. Mussels growing 44
in colander at
Ellwood Pier
3 a
e+ Il
Regression Equation pd
=-2.050+7.047X 2.395***
Y= 1.172+43.528X
Vol. 12; No. 4
THE VELIGER
Page 411
Table 13
Comparison of the frequency of check rings laid down by
Mytilus californianus when growing inside clumps at
Ellwood Pier with that frequency shown by the same
mussels growing freely on the outside of those clumps
The independent variable is shell length for the first group and
length of new growth for the second group (in centimeters). The
dependent variable is number of check rings.
Group
1. Original growth 240
within clumps
2. Final “free” growth 240
outside clumps
Regression Equation
Y=8.61+4.65X
1,476
2.972
1,476
Adjusted
Means at
n
vo
Bp
wn
x<—
14:09*** 21:97 205.73***
Y=1.7543.02X 10.42
Table 14
Comparison of the frequency of check rings laid down by
Mytilus californianus when growing inside mussel clumps
high up in the intertidal zone: (Top of mussel clumps
at Ellwood Pier, two samples), with that frequency
shown by mussels growing low in the intertidal (Inside
bottom of mussel clumps at Ellwood Pier, three samples)
The independent variable is shell length and the dependent variable
is the number of check rings.
#19
& 2 g 2 =
Be BES mR
Group Regression Equation nw g S v te
1. Top Pile 27 53 Y=3.2+6.7X 2505 33.468 IS) fe pdaratas
2. Top Pile 26 51 Y=1.5+6.8X 32.217
3. Bottom Pile 27 42 Y=9.8+3.9X 27.452
4. Bottom Pile 26 62 Y=7.9+4.7X 28.960
5. Bottom Pile 31 32 WS IIa FSep.< 27.393
F
Individual D. E Test on B 1,230
1.2./3.4.5 45.99***
Individual D. F. Tests on Adjusted Means
1.2./3.4.5 50.55***
mussels. Further to this, a positive correlation may be
said to exist between high check ring frequencies and
the action of any factor tending to disturb and inhibit
growth, i. e., such as found in mussels growing on exposed
shores, high up in the intertidal, or in the center of
clumps. Mussels in such situations show much higher
check ring frequencies than those taken from quiet areas
(Ellwood Pier), low positions within the intertidal or
on the outside of mussel clumps.
It would seem that 3 separate factors are involved in
the formation of check rings on Mytilus californianus:
1. There is some inherent ring-laying pattern which oper-
ates even in extreme shelter.
2. A mussel growing freely on the outside of a clump may
increase its frequency of ring formation in response to
periodic disturbances such as heavy waves, unduly low
tides, hot weather, etc.
3. Perhaps a mussel confined within a clump, and there-
fore growing extremely slowly (Harcer, 1967), may
occasionally experience “favorable” periods in which it is
able to grow a little. Such a period might be established
by movement of the mussel clump by wave action, so
allowing a particular mussel within the clump to be
relieved of pressure imposed on it by its neighbors for a
short time. Subsequent growth would then give rise to an
interval between rings. In this case the rings themselves
Page 412
THE VELIGER
Vol. 12; No. 4
60
50
40
30
a i
0.2 0.3 0.4 0.5 1.0 10.0
AA a a
Wave Force (kg — cm/cm?)
Adjusted Mean Check Rings
Figure 8
Log log plot of wave force against the mean number of check rings
on Mytilus californianus shells (adjusted to a shell length of 5 cm)
The © represents the mean ring frequency for a sample of mussels
taken from the outer pilings of a pier at Cayucos Beach, San Luis
Obispo County, California. The [_] represents the mean check ring
frequency for a sample taken from a mid tide rock platform at
Monterey Peninsula, Monterey County, California. Both these points
have been projected onto a line connecting the points indicating
check ring frequency obtained from mussels growing at known wave
impact values. The bar representing variation of check rings is
proportional to twice the standard error of the mean (on each side).
1. Stearns Wharf 2. outside end of Ellwood Pier
3. Ellwood Shore 4. Goleta Point
The line has been eye-fitted to the points
would probably be established during the time in which
longitudinal growth was prevented. In this connection it
should be noted that large mussel clumps, attached to
Ellwood Pier pilings, move visibly as waves thrust against
them; intermittent movement of this kind might allow
streams of water to pass through the clumps at some
times, but prevent this flow at others.
At least two causal mechanisms seem to be responsible
for high frequency ring formation: increased wave im-
pact etc (such as would be experienced by a mussel on
the outside of a clump) is one such process; the other
seems to be instigated as the result of confinement of
animals and the consequent uncertain growth within the
matrix of a clump. Such a mussel, so placed within the
clump, may experience little growth at infrequent inter-
vals (with each favorable interval providing the small
space between the previous ring and that established
when conditions favoring elongation are terminated).
Mussels establish rings in response to both factors,
but it is possible to distinguish individuals which have
developed inside a clump at a glance from those develop-
ing on the exterior. The projecting ridges of the rings in
the case of the former group tend to be sharply defined,
inverted V’s, in cross section, whereas the latter group
yield rings that tend to resemble inverted U’s, in cross
section.
Adjusted Mean Check Rings
20.30
40 .50
aa © @
A Wave Force (kg — cm/cm?)
Figure 9
Log log plot of wave force against mean number of check rings on
Mytilus edulis shells (adjusted to a shell length of 5 cm)
The bar representing check ring variation is proportional to twice
the standard error (on each side). Station 1 is a submerged electrode
cable at Ellwood Pier, 2 is Stearns Wharf, 3 is outside end of Ellwood
Pier, 4 the outer end of Cayucos Pier and 5 the inner region (surf
zone) of Cayucos Pier, San Louis Obispo County, California. The
exposure rating for the outer end of Cayucos Pier (Point A) was
obtained from the Mytilus californianus exposure plot (Figure 8).
The exposure value for the point labeled “inner end Cayucos (Point
5) was obtained by doubling the exposure value for the outer end of
this same pier, since the inner regions of Ellwood Pier are about
twice as exposed as the outer regions
The straight line was fitted by eye to the points
Vol. 12; No. 4
THE VELIGER
Page 413
Table 15
Comparison of the frequency of check rings laid down by
Mytilus californianus when growing freely high up in
the intertidal one (outside top of mussel clumps at
Ellwood Pier, two samples), with that frequency shown
by mussels growing low in the intertidal on the outside of
mussel clumps at Ellwood Pier (Three samples)
The independent variable is growth increment (from February 1966
to 22 June 1966) measured in centimeters and the dependent vari-
able is number of check rings.
1 © 2 be =)
N 5 q Sh A 395— we
Group Regression Equation ) oo || a
Lax
1. Top Pile 26 51 Y=2.942.5X 3.73N.S. 6.018 Wea heretes
2. Top Pile 27 53 NYG IP 2A axe 6.357
3. Bottom Pile 26 62 Y=2.342.7% 5.739
4. Bottom Pile 27. 42 Wess ipa 4.496
5. Bottom Pile 31 32 Y=1.04+3.5x 5.336
F
Individual D. F. Tests on Adjusted Means 1,230
1.2./3.4.5 23.0 2a
It seemed possible that the number of rings per unit
shell length could be used as an indication of the amount
of harassment mussels had been exposed to during their
growth, providing that consideration was lent only to
mussels from similar positions within the clumps. The
following observations lend further weight to this hypo-
thesis.
If mussels are taken from the outside of the clumps
from a fixed intertidal level, a correlation can be estab-
lished between the wave impact value (measured as re-
ported) to which they have been exposed and the fre-
quency of check rings per 5 cm individual (Figures 8, 9).
Such a relationship indicates that the check rings can be
used to estimate exposure at a particular place or, more
important, to provide an objective method for comparing
the exposure between geographical locations. A log log
scale was used to plot ring frequency against wave im-
pact value in Figures 8 and 9 because, as shown, the points
then fell on a straight line.
As mentioned previously, a lower limit was found to
exist for the frequency of check rings a mussel may lay
down, and it would seem that an upper limit might also
exist. This would be imposed by the width of the rings
themselves, i. e., only a fixed number could exist between
two points (if one assumes a minimum thickness for a
ring). To determine whether the frequency of check
rings on mussels can be used to estimate the wave impact
experienced by a particular region of shore line, I have
plotted in Figure 8 two points representing check ring
frequencies obtained from samples of mussels taken from
two places where I had not previously obtained wave
impact values. The locations were:
1. The outer pilings of a pier belonging to the Standard
Oil Company at Cayucos Beach, San Luis Obispo
County, California.
2. An intertidal rock platform at Monterey Peninsula,
California.
The first point, when projected onto the line connecting
the points indicating check ring frequency obtained from
mussel populations at known wave impact values, indi-
cates that this pier, located on a section of unprotected
coast line, experiences far greater exposure than that
experienced by Ellwood Pier.
The second point, I think, probably represents the max-
imum frequency of growth rings which can occur on a
5 cm mussel growing on the outside of a clump in ex-
posed conditions. By this I mean that, even if mussels
were grown in areas which received twice the wave shock
experienced by the Monterey mussels, they would be
physically incapable of laying down many more rings.
In both cases the check ring frequency yields values for
the exposure which these populations of mussels experi-
ence which is matched by subjective estimates.
A similar relationship between exposure and shell ring
frequency exists for Mytilus edulis (Figure 9), although
this is far less clear than the M. californianus example.
Although Barker (1964) has suggested that a positive
correlation exists between mean annual temperature and
the thickness of growth layers in 3 species of bivalves,
Mercenaria mercenaria, Mactra solidissima and Anadara
Page 414
ovalis, his data do not exclude the possibility that turbu-
lence or wave impact have played some part. However,
if this correlation reflects the action of temperature as a
causative modifying factor in the formation of growth
layers then one might expect that mussels growing in
colder waters would have more check rings than those
growing in warmer waters, all other things being equal.
SUMMARY anp CONCLUSIONS
The wave impact measuring device described provides
the intertidal ecologist with a simple method of estimat-
ing wave impact at various places. A biological recorder
in the form of mussel check ring frequency may also be
used where 2 or 3 comparisons are to be made within one
particular area. Since several factors are known to affect
the frequency of ring production, i.e., height on shore,
position within mussel clump, etc, great care must be
used when selecting the samples for analysis to ensure
that they are taken from comparable positions. I regard a
sample of around 50 individuals to be the minimum
necessary to establish the mean check ring frequency for
any one place. Furthermore, all sizes of mussels directly
exposed to wave impact should be equally represented
within this sample.
I have indicated that several trends in the morpholog-
ical characteristics of sea mussels can be linked with
changes in wave impact. In a later paper I will show
that an understanding of these trends was important in
investigating the nature of the competitive interaction
between Mytilus edulis and M. californianus on the coasts
of Southern California.
THE VELIGER
Vol. 12; No. 4
ACKNOWLEDGMENTS
This work forms part of a Ph. D. dissertation presented at
the University of California at Santa Barbara. I wish to
thank Dr. J. H. Connel, Dr. D. E. Landerberger and Dr.
J. Stimson for considerable advice and support. I wish
also to thank the Signal Oil and Gas Company for making
their premises at Ellwood available for ecological re-
search.
LITERATURE CITED
BALLANTINE, WILLIAM
1961. _A biologically defined exposure scale for the comparative
description of rocky shores. Field Studies 1 (3): 1-19
Barker, RicHarp M.
1964. Microtextural variation in pelecypod shells.
logia 2: 69 - 86
Bascom, WILLARD
1964. | Waves and beaches, the dynamics of the ocean surface.
Anchor Books, Doubleday « Co., Inc. Garden City, N. Y. 267 pp.
E1rion Jones, W. & ANDREAS DEMETROPOULOS
1965. A quantitative approach to the exposure problem.
Br. phycol. Bull. 2 (6): 516-517
Harcer, JoHN R. E.
1967. Population studies on Mytilus communities.
dissertation, Univ. Calif. at Santa Barbara
Li, Jerome C. R.
1964. Statistical inference (I).
Arbor, Mich., 658 pp.
Moore, Hitary Brooke
1935. The biology of Balanus balanoides. Part IV - Relation
to environmental factors. Journ. Marine Biol. Assoc. U. K.
20: 279 - 307
SouTHWappD, A. J. & J. H. Orton
1954. The effects of wave action on the distribution and
numbers of the commoner plants and animals living on the
Plymouth breakwater. Journ. Marine Biol. Assoc. U.K.
33: 1-19
Malaco-
Ph. D.
Edwards Bros., Inc., Ann
Vol. 12; No. 4
PERVEWIGER
Page 415
Supplementary Comments on Deep Water Volutidae
from the South China Sea and South Africa
BY
HARALD A. REHDER
Division of Mollusks, Smithsonian Institution, Washington, D. C. 20560
(Plate 61)
ADDITIONAL MATERIAL and new information kindly fur-
nished me by correspondents have impelled me to publish
this paper as a supplement to two earlier papers of mine
(ReHveER, 1967; 1969) dealing with deep-water members
of the family Volutidae from the South China Sea and
South Africa.
For cooperation in making this contribution possible I
thank the following individuals: Mr. & Mrs. Francis S.
Harmon of New York City, Mr. William E. Old of the
American Museum of Natural History, Mr. John E.
duPont, Director of the Delaware Museum of Natural
History, and Mr. Clifton S. Weaver of Honolulu, Hawaii.
As in my earlier papers I have used the initials DMNH
for the Delaware Museum of Natural History and USNM
for United States National Museum (now more properly
known as the National Museum of Natural History).
Sigaluta pratasensis REHDER, 1967
(Plate 61, Figure 2)
1967. Sigaluta pratasensis REHDER, Pacific Science 21: 182;
figs. 1 - 4
The receipt for examination of a large example of this
species has induced me to place a note concerning it on
record. This specimen was obtained by Mr. and Mrs.
Francis S. Harmon from a fisherman in Kaohsiung, Tai-
wan, who is said to have obtained it from moderately deep
water southwest of Taiwan. It was collected, therefore,
somewhere in the South China Sea, probably between
Taiwan and the Pratas Reef, which is the type locality of
the species.
The specimen, now in the collection of Mr. and Mrs.
Francis S. Harmon, consists of 44 whorls and has the fol-
lowing measurements: length 103.5mm, width 53 mm,
length of aperture 73.5 mm. The surface is dull, not glossy
as in the young specimens on which the original descrip-
tion was based, and shows considerable etching by marine
organisms (probably cleaned off by the collector). The
color of the shell has a more grayish tone than in the
holotype and paratype, being a light grayish yellowish
brown (no. 79) to brownish pink (no. 33) (ISCC-NBS
color names, KELLY & Jupp, 1965). The columella is
brownish with two spirally ascending plaits which are of
a pale pinkish white color.
It is hoped that further examples of this species, which
we now know reaches a length of at least 4 inches, will be
found in the area between Taiwan, South China and the
northern Philippines, and that the fishermen may be
encouraged to preserve the soft parts so that its anatomy
may be studied.
Fuswoluta capensis (THIELE, 1925)
(Plate 61, Figures 4, 6)
1925. Glypteuthria (?) capensis THEE, Wiss. Ergebn. deutsch.
Tiefsee-Exp. “Valdivia”, 1898-99, 17 (2): 179; plt. 19,
fig. 27
1959. Fusivoluta capensis (THIELE), BARNARD, Ann. South
Afr. Mus. 45: 30 - 31 (in part)
I have been able to examine two color transparencies of
the type of THIELE’s species, taken by Mr. Hans Lehman
of Black Star. From these slides, which are reproduced in
black and white on Plate 61, Figures 4 and 6, I have con-
structed a description of this species to supplement the
brief four-line description given by THIELE.
Description: Shell small, ivory-white in color, obovately
fusiform in shape, suture deeply impressed, almost chan-
neled, whorls of spire rather flattened, body whorl slightly
convex. Protoconch of about 2+ (?) nuclear whorls,
smooth, bulbous. Postnuclear whorls strongly axially
ribbed, the ribs retractively curved, about 22 - 23 on last
whorl. The ribs are crossed by fine spiral cords, about 10 in
number on penultimate whorl, the subsutural cord pro-
nounced, giving the suture an almost channeled appear-
ance. Aperture narrowly ovate, the anterior canal rather
broad at the slightly constricted base.
Measurements (from THIELE): Height, 11 mm; width,
4.5 mm.
Remarks: In my paper on South African deepwater vo-
lutes (REHDER, 1969, p. 205) I followed BaRNarp (1959,
p. 30) in synonymizing Glypeuthria capensis TOMLIN
Page 416
THE VELIGER
Vol. 12; No. 4
(1932, p. 165) with THreLe’s species. An examination of
a photograph of the type of THIELE’s species, however,
leads me to believe that TomMLin’s species, renamed sculp-
turata Tomun (1945, p. 135), should, at least for the
time being, be considered a distinct species.
From Fusivoluta sculpturata (Tomun) FE capensis dif-
fers notably in its size. The type, consisting of 6 whorls
(THIELE says 53 whorls, but the photograph shows 6 or
even slightly more whorls), measures only 11 mm, while
the type of FE sculpturata, which ToMuIn states consisted
of 7 whorls (though here again the photograph seems to
show a shell of 64 whorls), measures 29 mm, almost 3
times as large; other specimens of F sculpturata that are
even somewhat larger have only 6 to 64 whorls. Thus,
specimens of F sculpturata with approximately the same
number of whorls possessed by the type of F capensis are
almost 3 times as large as the latter. The spire of F cap-
ensis THIELE is not as slender as in F sculpturata, the
whorls appear to be channeled, the axial ribs are more
numerous (22 - 23 on the last whorl, instead of 16 - 18 as
in EF sculpturata), the spiral cords are subequal rather
than unequal, and there is a strong subsutural cord present
instead of the series of 5 fine cords below the suture.
Fusivoluta sculpturata (TOMLIN, 1945)
(Plate 61, Figures 1, 3, and 5)
1932. Glypteuthria capensis Tomuin, Ann. South Afr. Mus.
30: 165; fig. 6 (not THIELE, 1925)
1945. Glypteuthria sculpturata ToMLIN, Journ. Conch. 22: 135
1957. Fusivoluta capensis (THIELE), BARNARD, Journ. Conch.
24: 210 (not Tuer, 1925)
1959. Fusivoluta capensis (THIELE), BARNARD, Ann. South
Afr. Mus. 45: 30 - 31 (in part)
1965. Fusivoluta capensis (THIELE), WEAVER, Hawai. Shell
News 13 (3): 7; figs. 5, 6
1969. Fusivoluta capensis (TuHtELe), REHDER, The Veliger
11: 205; plt. 42, fig. 32 (in part)
Description: Shell elongate, obovate, with rather attenu-
ate spire, grayish white, whorls moderately convex, ap-
pressed at suture. Protoconch consisting of probably 2
(corroded) nuclear whorls; postnuclear whorls slightly
shouldered and gently convex, bearing arcuate axial ribs
numbering 16 to 18 on last whorl. The ribs are crossed by
spiral cords sometimes varying in strength and crowded,
or subequal and more or less separated; between the slight
shoulder and the appressed suture the spiral cords are
noticeably finer. Aperture ovate, last whorl usually not
constricted at the short broad anterior canal. Operculum
ovate-unguiculate, nucleus terminal.
Measurements (in mm)
Height Width
Holotype (from Tomtin) 29.0 12.0
S.A. M. (from Barnarp, 1959) 38.0 15.0
DMNH 10668 30.0 13.0
DMNH 10123 742) ff 11.9
Remarks: The differences between this species and F
capensis THIELE have already been noted above. I am as-
signing with some doubt the specimens in the Delaware
Museum of Natural History, which I have examined, to
ToMLIn’s species, which I know only from the original
figure and the descriptions by Tomuin and Barnarp (1959,
p. 31). The outer lip on both the Delaware specimens is
imperfect, and they seem to have suffered injury of some
kind; neither shows the evenly and broadly arcuate out-
line and lack of constriction at the anterior canal shown
in ToMLin’s original figure.
In sculptural characters, however, the specimens in the
Delaware Museum of Natural History agree with Tom-
LIN’s type, with the exception that the nature of the spiral
cords on the body whorl is variable; in one specimen (DM
NH 10123) the cords are crowded and rather unequal as
in the holotype; in the other (DMNH 10668) the cords
are distant and equal. However, until further material is
available and the types can be reexamined, it is better to
unite all these specimens under one name.
LITERATURE CITED
BarNARD, KEPPEL Harcourt
1957. Glypteuthria capensis: a generic correction.
Conch. 24: 210
1959. Contributions to the knowledge of South African marine
Mollusca, Part II :Gastropoda: Prosobranchiata: Rhachiglossa.
Ann. South Afric. Mus. 45 (1): 1-237; 52 figs. (June)
Ketty, KENNETH Low & DEANE BREWSTER JUDD
1965. The ISCC-NBS method of designating colors and a
dictionary of color names. Nat. Bur. Standards Circ. 553
(reprint ed.) v+158 pp. ISCC-NBS color-name charts illus-
trated with centroid colors (Suppl. to NBS Circ. 553)
REHDER, HarALD ALFRED
1967. A new genus and two new species in the families Voluti-
dae and Turbinellidae (Mollusca: Gastropoda) from the West-
ern Pacific. Pacif. Sci. 21: 182 - 187; 11 figs.
1969. Volutocorbis and Fusivoluta, two genera of deepwater
Volutidae from South Africa. The Veliger 11 (3): 200
to 209; plts. 40 - 43 (1 January 1969)
THIELE, JOHANNES
1925. | Gastropoda der deutschen Tiefsee-Expedition. II. Teil.
Wiss. Ergebn. Deutsch. Tiefsee-Exped. “Valdivia,’ 1898-1899.
17 (2): 35-382; plts. 13-46; 31 text figs.
ToMLIN, JoHN Reap LE BRocKTON :
1932. Reports of the marine Mollusca in the collection of the
South African Museum. VI - VIII. Ann. South. Afr. Mus.
30: 157 - 169; 10 text figs.
1945. ‘Two South African species renamed.
22: 135
WEAVER, CLIFTON STOKES
1965. _ Volute problems.
8 text figs.
Journ.
Journ. Conch.
Hawaiian Shell News 13 (3): 7;
lo
THE VELIGER, Vol. 12, No. 4 [REHDER] Plate 61
Figure 1 X a —_ Figure 3
Figure 4 Figure 5 Figure 6
Figures, 1 and 3: Fusivoluta sculpturata (ToMLIN)
DMNH 10668 (xX 2-)
Figure 2: Sigaluta pratasensis REHDER
ES. Harmon Coll. (X 1-)
Figures 4 and 6: Fusivoluta capensis (THIELE)
Holotype, Zool. Mus. Berlin (X 6)
Figure 5: Fuswoluta sculpturata (Tomuin)
DMNH 10123 (X ‘1.5-)
Vol. 12; No. 4
THE VELIGER
Page 417
Uptake of Sea Water into the Fluid Spaces
of the Prosobranch Gastropod, Acmaea scutum '
BY
H. H. WEBBER’
Department of Zoology, University of British Columbia, Vancouver 8, Canada
INTRODUCTION
THE CIRCULATORY SYSTEM of most gastropod mollusks
functions in part as a hydrostatic skeleton. In the past it
was believed that gastropods could incorporate sea water
into the blood space to facilitate the expansion of the
foot. However, the current view is that, with one excep-
tion (the family Naticidae) a constant blood volume in
the circulatory system is sufficient to cause the expansion
of the foot of gastropods (Morris, 1950; CHapMaN,
1958; Brown & TuRNER, 1962; Brown, 1964; BERNARD,
1968; RusseELt-HunNTER & ApLey, 1969). For the Nati-
cidae Morris, op. cit., BERNARD, op. cit., and RUSSELL-
HunTER & APLEy, op. cit., have described a system of
aquiferous ducts that are separate from the blood system
and fill with sea water during foot expansion.
This study shows that the limpet Acmaea scutum
RaTHKE, 1833, which does not use the hydrostatic skele-
ton to expand the foot in the manner of most gastropods,
does however, demonstrate large changes in total volume.
Evidence is presented showing that this change in vol-
ume results in part from the incorporation of large quan-
tities of sea water directly into the blood space.
MATERIAL ann METHODS
Most limpets used were collected from Jordon River,
British Columbia. Laboratory maintenance is described in
WEBBER & DEHNEL (1968). Acmaea scutum used in
methylene blue experiments were collected at Stillwater
Cove, Monterey, California.
1 This study was aided by grants from the National Research
Council of Canada and the National Science Foundation of the
United States to Dr. Paul A. Dehnel. I thank Dr. J. H. Phillips,
Director of Hopkins Marine Station for generously providing
equipment and facilities for part of this study.
2 Present address: Department of Biology, Wake Forest Univer-
sity, Winston-Salem, North Carolina 27109
Measurements of Volume Change
Animals were maintained in sea water for 24 hours
before use. The method of volume measurements was as
follows. The animal was taken from the experimental
salinity and the foot was compressed gently with absorbent
tissue. Volume was determined by removing the limpet
from sea water, compressing the foot gently with absor-
bent tissue, and weighing the volume of sea water dis-
placed by the animal. The method was accurate to +0.1
ml. ‘To measure changes in volume, limpets were returned
to sea water with the dorsal surface of the shell against
substrate so the animal could not attach with the foot.
Animals were maintained in this position for the desired
time period. The volume of the animal was again deter-
mined after gently shaking to remove water from the
nuchal cavity and space between the foot and shell. The
increase in volume was determined by subtraction. The
volume of the soft body parts alone was determined by
estimating the volume of the shell separately and sub-
tracting this value from the total volume.
Blood Amaranth Samples
Samples for the determination of blood amaranth values
were taken from the visceral sinus through an incision
in the foot muscle. Approximately 100m liter samples
were measured colorimetrically without dilution at 520
mpl, using sea water as a blank.
Blood Inulin Samples
Blood samples for the measurement of inulin were taken
from an incision through the foot into the visceral sinus.
Fifty microliter aliquots of blood were assayed for inulin
by the anthrone method of Younc & Ratsz (1952).
RESULTS
When Acmaea scutum is removed from the substrate and
placed in sea water upside down so it cannot right itself,
Page 418
THE VELIGER
Vol. 12; No. 4
over a period of time the soft parts of the limpet appear
to swell. The results recorded below document and char-
acterize this change in size.
In measuring the volume increase, it was important
that limpets at the start of an experiment had the same
relative water content. Before measuring the starting
volume the animal was blotted with absorbent tissue to
remove excess water. A test using regression slopes (Table
1) showed that all limpets had the same relative water
content when the starting volume was measured.
Table 1
Regression equations of initial water content plotted
against dry weight for Acmaea scutum used in experiments
on uptake of water at constant salinity. The p value
indicates the probability level at which the regression slope
is significant
average water regression equation P
content
NC = eS SP Ds. 0.01
83.7% (n = 60) Y = 0.81 + 4.32X 0.01
Y = 0.68 + 4.50X 0.01
The change in volume with time is given in Table 2.
The volume of the animals increased even though the
salinity of the water was constant. By 6-12 hours in
this inverted position, limpets increased the volume of the
soft parts by approximately 100%. The variation indi-
cated that only large differences could be considered sig-
nificant. The purpose of the experiment was to demon-
strate that Acmaea scutum could show an increase in
volume of the soft body parts when held at a constant
salinity.
Nature of the Volume Change
The blood and urine of Acmaea scutum are essentially
sea water. The blood and urine concentrations of the
major ions (Na*, Cl, Mg**, and Ca‘) are the same as
the corresponding concentrations in sea water (WEBBER
& DEHNEL, 1968). Ion values for blood and urine samples
taken at all stages during increase in volume remained
constant, i. e., were the same as sea water. The observed
increase in volume, then, must have been achieved by
taking in both ions and water.
Uptake of Amaranth
To determine if the water uptake response involved all
fluid spaces of the limpet, the red dye amaranth was
Table 2
Increase in volume of soft body parts of Acmaea scutum
in 100% sea water. Part A is the number of animals of a
sample of 10 showing an increase in soft body parts of 5%.
Part B is the mean increase in volume (per cent) for those
animals showing an increase of 5%. Part C is the blood
concentration of amaranth (mg/l) at a concentration of
0.025 mg/l
Part A
number of animals out
of 10 showing increase
in volume
Time in hours
[pe SPO Ge ee ES
2 0. 50, 50 10
Part B
means of increase in volume
(per cent)
Time in hours
14 3 6 12 24 48
70.7 71.7 86.2 98.2 100.3 100.5
Part C
Blood concentration of amaranth
Time in hours
14 3 6 12 24 48
dye conc. 0.013 0.012 0.012 0.011 0.014 0.010
n 4 7 7 8 7 9
added to the sea water to act as a tracer. Comparison
(t-tests) of results of volume changes in sea water with
and without amaranth showed the presence of the dye
did not significantly (p=0.01) alter the increase in
volume. When limpets increased in volume in “amaranth
sea water” the dye became distributed throughout the
tissues. All parts of the animal “blushed” — including the
gill and mantle fringe.
After volume changes, the values of amaranth in the
blood were determined. There is a relationship between
blood amaranth values and volume increase (Table 2).
The average volume increase was around 100%, indicat-
ing the volume of water taken up was equal to the
starting volume of the soft parts. The average blood dye
value for these animals (0.012g/) was around $ that of
sea water (0.025g/1). It appears the increase in volume of
soft parts was facilitated by bulk movement of sea water
Vol. 12; No. 4
THE VELIGER
Page 419
with the dissolved dye into the fluid spaces of the limpet.
The final blood dye concentration (4 that of sea water)
would then be due to the diluting effect of the fluid
of the starting volume. It is not possible to analyze more
critically the relationship between water and dye uptake
because (1) the intracellular distribution of amaranth,
if any, was not known, and (2) the extent of the blood
space at any given time was not known. Amaranth did
enter the kidney space as well as the blood space. Animals
stimulated to contract after volume increase in amaranth
sea water would evacuate urine that was red from the dye.
Uptake of Inulin
Inulin is generally believed not to enter cells (WHITE et
al. 1959), although Scorr et al. (1964) report the ab-
sorption of inulin from the proximal tubule of Necturus
kidney. To determine if Acmaea scutum could also take
up inulin during volume increases limpets were placed in
sea water solutions of inulin. Table 3 gives the results.
Again limpets increased in volume around 100%. The
average blood inulin value was 2.8g/1 compared with
4¢/] in the surrounding sea water. If the starting volume
was considered as a water space the doubling of volume
by uptake of water and 4¢g/1 inulin would result in a final
blood inulin value of 2.0g/1. However, since inulin prob-
ably did not penetrate intracellularly, the fluid available
for dilution would be less and the observed blood inulin
value of 2.8g/1 would be expected. As with amaranth,
the inulin data support the hypothesis that the increase
in volume occurred by a bulk movement of sea water into
the blood space.
Table 3
Blood inulin concentration (g/l) after Acmaea scutum
had shown an increase in volume of soft body parts by
taking up sea water. Each experimental salinity contained
an inulin concentration of 4g/I1
average average
% increase n inulin conc. n
113.8 30 2.85 29
Mechanism of Volume Increase
Histological sectioning showed no pores leading into the
blood space. ‘To determine if water was passing into the
radula sac or intestine the volume increase was followed
in a sea water solution of the vital dye methylene blue.
During the volume increase the gut but not the
radula sac stained blue. To insure that it was not a
case of selective staining, 1 cc of 0.1% methylene blue
was injected into the blood space of each of 5
limpets. After 2 hours, the gut and radula sac were ex-
amined. Both were equally but lightly stained (2 out of
a scale of 4). Moreover, when the radula and radula sac
were removed from 5 limpets and stained in vitro in 0.1%
methylene blue for 15 minutes, the radula sac was stained
(3 out of a scale of 4). These data support the idea that
sea water was passed into the gut. No direct evidence is
available, however, that sea water passes from the gut
into the blood space.
DISCUSSION
In this study it is shown that the limpet Acmaea scutum
was capable of taking large quantities of sea water into
both the blood and urine spaces. Limpets could take up a
volume of sea water approximately equal to the starting
volume of the soft parts. It was not possible to separately
measure the extent of the increase of blood and urine
spaces. Data using the vital dye methylene blue suggest
that sea water passes into the blood space through the gut.
This response of Acmaea scutum differs from the in-
crease in volume shown by the Naticidae (moonsnails).
There, the large volume increase is due to uptake of sea
water into “aquiferous ducts” that are separate from the
blood space (Morris, 1950; BERNARD, 1968; RussELt-
Hunter & ApLey, 1969). In A. scutum the sea water
taken in during volume increase, in part at least, mixes
directly with the blood.
It is difficult to explain how the change of volume of
Acmaea scutum is connected to the function of the hydro-
static skeletons. Limpets cannot withdraw into their shell,
so the uptake of sea water is probably not normally used
for foot expansion. These experiments were performed
with the limpets turned upside down. However, it is
unlikely that the uptake of sea water into the fluid spaces
is unique to the animal being in this position. Possibly, sea
water uptake is an adaptation to intertidal exposure. The
importance of free water in the mantle cavity and pallial
groove (extravisceral water) of limpets in decreasing
desiccation effects have been shown by SEGAL & DEHNEL
(1962). When they removed the extravisceral water from
Acmaea limatula CARPENTER, 1864, the limpet showed a
more rapid increase in the total osmotic pressure of the
blood when desiccated. As well, SHotwett (1950)
Page 420
showed that animals having the greatest exposure time
have the largest extravisceral water space. In Acmaea
scutum the ability to take sea water into the circulatory
system could be an important adaptation to desiccation
stress in that it results in a larger fluid volume to act as
an osmotic buffer.
SUMMARY
1. Evidence is presented to show that Acmaea scutum
could increase the water content of the soft body parts at
a constant salinity. The increase in water content resulted
from sea water entering the fluid spaces from the external
environment.
2. While taking up sea water from the external environ-
ment, the molecules inulin and amaranth could also pass
into the blood space from the external sea water when
these molecules were dissolved in experimental salinities.
3. Using methylene blue as a tracer, it appears that sea
water enters the fluid spaces of the limpet through the
gut.
LITERATURE CITED
BERNARD, E.R.
1968. The aquiferous system of Polinices lewisi (Gastropoda,
Prosobranchiata) . Journ. Fish. Res. Brd. Canada 25:
541 - 546
THE VELIGER
Vol. 12; No. 4
Brown, A.C.
1964. Blood volumes, blood distribution and sea-water spaces
in relation to expansion and retraction of the foot in Bullia
(Gastropoda). Journ. Exp. Biol. 41: 837 - 854
Brown, A.C. « L. G. W. TuRNER
1962. | Expansion of the foot in Bullia (Gastropoda) . Na-
ture 195: 98 - 99
CHAPMAN, GaRTH
1958. The hydrostatic skeleton in the invertebrates. Biol.
Rev. 33: 338 - 371
Morris, Murret C.
1950. Dilation of the foot in Uber (Polinices) strangei (Mol-
luscs, class Gastropoda) . Proc. Linn. Soc. New South Wales
75: 70 - 80
RussELL-Hunter, W. D. & M. L. APLEY
1969. Pedal expansion in the naticid snails. II. Labelling ex-
periments using inulin. Biol. Bull. 135: 553 - 573
Scott, W. N., D. L. Maune, I. SHEHADEH « A. K. SoLoMON
1964. Inulin and albumin absorption from the proximal tub-
ule in Necturus kidney. Science 146: 1588 - 1590
SecaL, Eart & Pau, Aucustus DEHNEL
1962. Osmotic behavior in an intertidal limpet, Acmaea lima-
tula. Biol. Bull. 122 (3) : 417 - 430; 5 figs.; 1 table
SHOTWELL, JESSE ARNOLD
1950. Distribution of volume and relative linear measurement
changes in Acmaea, the limpet. Ecology 31: 51 - 61
WEBBER, HERBERT H. « PauL Aucustus DEHNEL
1968. Ion balance in the prosobranch gastropod Acmaea
scutum. Comp. Biochem. Physiol. 25: 49 - 64
Wulire, A., P Hanpier & E. L. Smiru
1959. _— Principles of Biochemistry.
1106 pp.
Youn, M. « L. Ratsz
1952. Anthrone procedure for determination of inulin in bio-
logical fluids. Proc. Soc. Exp. Biol. Med. 80: 771 - 774
McGraw-Hill, New York,
Vol. 12; No. 4
THE VELIGER
Page 421
Notes on the Deep Water Calliostomas of the Panamic Province
with Descriptions of Six New Species
JAMES H. McLEAN
Los Angeles County Museum of Natural History
goo Exposition Boulevard, Los Angeles, California 90007
(Plate 62)
Tue PaNnamic specigs of the trochid genus Calliostoma
may be grouped by depth of occurrence: one group of
species occurs at low tide in rocky sublittoral zones, and
on shallow offshore bottoms to depths of 20 fathoms,
while another group of species is dredged only at depths
of 30 fathoms and deeper. Only 2 deep water species
have previously been known in the Panamic province. A
third species known from the Peruvian province is added
to the list and 6 new species are here described, based on a
relatively small amount of material. In comparison to
the numerous species of Calliostoma in the tropical west-
ern Atlantic (CLENcH & TurRNER, 1960), the number of
species in the eastern Pacific is relatively sparse. The
comparative lack of extensive deep dredging in the east-
ern Pacific may account for the paucity of known species.
A more nearly equivalent number of species may even-
tually be known.
Sources of the new species described herein are as fol-
lows: ‘The Templeton Crocker Expedition of 1936 pro-
duced one of the species. Three result from the Allan
Hancock Pacific Expeditions and are part of the Allan
Hancock Pacific Foundation Collection, now on loan to
the Los Angeles County Museum of Natural History. Two
result from recent dredging at Santa Cruz Island, Gala-
pagos Islands, by André and Jacqueline DeRoy, the orig-
inal specimens of which were kindly forwarded to me by
Mr. Anthony D’Attilio of the San Diego Natural History
Museum. I am grateful to the institutions and collectors
mentioned for the opportunity to work with this material.
Abbreviations for institutions cited in the text are as
follows:
AHF Allan Hancock Foundation (collection on loan
to LACM)
CAS California Academy of Sciences, San Fran-
cisco
LACM Los Angeles County Museum of Natural
History
SDNHM_ San Diego Natural History Museum
SSB S. Stillman Berry Collection, Redlands,
California
SU Stanford University Collection, Stanford,
California
USNM United States National Museum
No attempt has been made at this time to assign any
of the eastern Pacific species of Calliostoma to subgenera.
CLENCH & TuRNER (1960) recognized three subgeneric
groups based on radular and jaw morphology. In estab-
lishing new subgeneric taxa they failed to consider other
generic units in the subfamily Calliostomatinae, many of
which have been used in a full generic sense by workers
in other parts of the world. My own radular studies on
the easern Pacific species are in progress.
Three of the 8 species treated here are recorded from
San Jaime Bank off Cape San Lucas and a 4" from the
Gorda Banks in the same vicinity. Further dredging at
these banks should prove productive.
Calliostoma iridium Daut, 1896
(Plate 62, Figure 1)
Calliostoma iridium Dat, 1896, p. 7; — 1902 p. 552,
plt. 39, fig. 3; — 1908, p. 348, plt. 19, fig. 5
Diagnosis: Whorls flat sided, spiral cording consisting of
2 strong, beaded cords at the periphery and one below the
suture, the area between these nearly smooth on the early
whorls but with 7 finely beaded cords increasing in promi-
nence on later whorls; base imperforate, sculptured with
numerous, weakly beaded spiral threads. Color, yellowish
pink with radiating flammules, with bronze pink irides-
Page 422
cence strongest on the early whorls. Height, 21.5, diameter,
19.2 mm (holotype).
Type Material: Holotype, USNM 122957; 7 paratypes,
USNM 122957a; 1 paratype, SSB 16962. Type locality:
USFC station 3387, Gulf of Panama, 127 fms.
Distribution: Gulf of Panama. In addition to the type
lot consisting of 9 specimens, there is 1 specimen, USNM
122956, USFC sta. 3391, Gulf of Panama, 153 fms.
Discussion: Calliostoma iridium may be distinguished
from other eastern Pacific species by the near absence of
sculpture between the subsutural and the 2 peripheral
cords on the early whorls.
Calliostoma nepheloide DALL, 1913
(Plate 62, Figures 2 to 5)
Calliostoma nepheloide Daur, 1913, p. 592; — 1925,
p. 9, pit. 24, figs. 2-3; — Keen, 1958, p. 256,
fig. 46 (upper figure only, copy DALL) ; — Par-
KER, 1964, p. 151
Diagnosis: Whorls flat sided, final whorl slightly con-
vex, spiral cording consisting of a strong, projecting,
beaded peripheral cord and about 12 fine beaded cords
between it and the suture on the final whorl. Base imper-
forate, defined by a broad, unbeaded cord below the peri-
pheral cord, base with about 18 weakly beaded cords,
more broadly spaced near the columella. Color, olive
green with darker flammules, basal cords with alternating
light and dark markings. Height, 25, diameter, 22 mm
(holotype).
Type Material: Holotype, USNM 96637. Type locali-
ty: USFC sta. 2804, Panama Bay, 47 fms. The holotype
has the lip broken back about 4 of a whorl. Dati’s 1925
illustration reconstructed the position of the lip.
Distribution: Point Abreojos to Cape San Lucas, Baja
California; Mazatlan, Sinaloa, Mexico to Gulf of Pana-
ma. Records: AHF 1711-49, 30 mi. S. Pt. Abreojos, Baja
California, 52 fms.; AHF 618-37, San Jaime Bank, off
Cape San Lucas, 75 fms. (Plate 62, Figure 5) ; off Mazat-
lan, Sinaloa, Mexico, 88 - 92 fms. (ParKER, 1964) ; AHF
273-34, Tenacatita Bay, Jalisco, Mexico, 45 fms.; SDN
HM 39799, off Manzanillo, Colima, Mexico, 52 fms.
(Plate 62, Figures 3, 4); LACM, Gulf of Tehuantepec,
Chiapas, Mexico, leg. D. Shasky, 45 fms.; CAS 17986,
Gulf of Chirqui, Panama, 35-40 fms. The species is
as yet unknown from the Gulf of California.
Remarks: The 2 immature specimens from San Jaime
Bank off Cape San Lucas differ from the others in having
THE VELIGER
Vol. 12; No. 4
the immediate subsutural cord more strongly beaded than
the others. This is the only significant feature of varia-
tion in the material examined. The olive coloration of
the species is distinctive.
Calliostoma fonku (PxHiuiprt, 1860)
(Plate 62, Figures 6, 7)
Trochus fonkit Puruiprt, 1860, p. 185, plt. 7, fig. 22
Calliostoma fonkii (Putt.) — Pirssry, 1889, vol. 11,
p. 371, plt. 57, fig. 48 (copy Puitiprr). — DALL,
1909, p. 240.
Diagnosis: Whorls flat sided, final whorl slightly con-
vex; spiral cording consisting of a smooth peripheral cord,
the lower part of which is covered by succeeding whorls,
and 3 prominent cords per whorl, the 2 uppermost coarse-
ly beaded, narrow raised ridges between major cords ap-
pearing on the final whorls; base imperforate, with up to
11 evenly spaced unbeaded cords. Color tan, with darker
cording. Height, 17.5, diameter, 16.2 mm (AHF 802-38).
Type Material: Holotype, not located. Type locality
uncertain: “Between Chiloé and the mainland of Peru”
(Pitssry, 1889).
Distribution: Galapagos Islands; Peru south to Chiloé
Island, Chile (43° S). Records: AHF 802-38, NW of
Charles (Santa Maria) Island, Galapagos Islands, Ecua-
dor (1°09’ S, 90°35’ W), 250 fms. (Plate 62, Figure 6) ;
AHF 371-35, Independencia Bay, Peru (14°15’S), 5
fms. (Plate 62, Figure 7). The two specimens here re-
ported are believed to be the first specimens known since
the original description.
Remarks: Although the 2 specimens are from widely
separated localities and vastly different depths, no essen-
tial points of difference are apparent. The Galapagan
specimen is colorless and the nacre is slightly leached,
while the smaller Peruvian specimen was live-taken and
is tan with brownish ribs.
Little can be said of the distribution and occurrence of
this species until more material is known. It may perhaps
not be a characteristically deep water species, judging at
least from its shallow occurrence in Peru.
Calliostoma gordanum McLean, spec. nov.
(Plate 62, Figures 8 to 10)
“Calliostoma nepheloide Datu.” — Kren, 1958, fig.
46 (lower figure only).
Diagnosis: Whorls slightly convex, spiral cording con-
Vol. 12; No. 4
THE VELIGER
Page 423
sisting of 2 peripheral cords with a finer intercalary cord
between; 7-8 strong beaded cords between sutures on
final whorl, secondary threading between cords on final
whorl; base imperforate, with 17 - 20 beaded cords. Col-
or whitish, with radiating light tan maculations, base un-
marked. Height, 19.6, diameter, 20.0 mm (holotype).
Description of Holotype: Shell of moderate size for the
genus, light, rather fragile. Postnuclear whorls 7, slightly
convex. Periphery rounded but defined by 2 somewhat
stronger spiral cords with a finer intercalary cord be-
tween. Fifth postnuclear whorl with a strongly beaded
subsutural cord and 6 thin, raised, unbeaded cords be-
tween it and the upper peripheral cord. By the 5" whorl
the intermediate cords are finely beaded and fine inter-
calary threads have appeared. By the 7" and final whorl
the intercalary threads have increased in size and are
beaded as are the primary cords. Base imperforate with 17
cords, intercalary threads between the outermost cords,
those nearer the columella more broadly spaced and bead-
ed in a radiating pattern. Color whitish tan with brownish
flammules, aperture iridescent with pink and green, base
unmarked.
Type Material: Holotype, CAS 13271; 3 paratypes, CAS
13272; 1 paratype, LACM 1268; 1 paratype, USNM
679551; 3 paratypes, SDNHM 40041; 1 paratype, SU
9985. Type locality: Gorda Banks, off southeastern Baja
California, CAS locality 17752. According to Dr. Leo G.
Hertlein, this station represented 26 hauls made by Croc-
ker and Beebe on the Templeton Crocker Expedition of
1936 in the vicinity of the Gorda Banks, approximately
23°02’ N, 109°31’ W, probable depth, 70 fms. Seven spe-
cimens, originally identified by A. M. Strong as Callio-
stoma nepheloide, were in this lot. Three additional spe-
cimens, evidently from the same lot, were found in the
Strong collection at the San Diego Museum.
Referred Material: One additional lot is known: AHF
531-36, San Francisquito Bay, Baja California (28°26’ N,
112°53’30” W), 10 fms., 2 specimens (Plate 62, Figure
10). The illustrated specimen from this lot is 13 mm in
height and has slightly more prominent spiral cording
but is otherwise typical. The relatively shallow depth
record of 10 fathoms for this lot may be anomalous.
Dredging off the tip of Baja California should produce
additional material of this species.
Discussion: Calliostoma gordanum may be separated
from C. nepheloide by its light color, its near absence of
spiral markings on the basal cords, and in having 2
rather than 1 major peripheral cords.
Calliostoma sanjaimense McLean, spec. nov.
(Plate 62, Figure 11)
Diagnosis: Whorls flat sided, spiral cording consisting of
2 strong, beaded peripheral cords, a strong subsutural
cord, and 5 strongly beaded intermediate cords; base
imperforate, with 11 broad, nearly smooth spiral cords.
Color yellow brown with light and darker markings par-
ticularly on the peripheral cords. Height, 20.0, diameter,
18.4mm (holotype).
Description of Holotype: Shell of moderate size, sturdy,
flat sided. Postnuclear whorls 7. Periphery angulate, de-
fined by 2 prominent beaded cords with a narrow inter-
calary cord between. A strong, beaded subsutural cord
is prominent on later whorls. On the 3 whorl there are
3 cords of equal strength; other cords arise as intercalary
threads that eventually become beaded and assume full
size, until in the final whorl there are 5 beaded cords and
4 intercalary threads between the subsutural and the
uppermost peripheral cord. Base imperforate, with 17
broad, low cords, interspaces of nearly equal width, the 3
cords close to the columella faintly beaded. Color yellow
brown with light and darker maculations especially on
the peripheral cords.
Type Material: Holotype, LACM-AHF 1269; 2 para-
types, LACM-AHF 1270, 1 paratype, USNM 679552.
Type Locality: San Jaime Bank, west of Cape San
Lucas, Baja California, Mexico, 22°50’30” N, 110°15’
W, 75 fms., Velero III station 618-37, 3 March 1937. The
3 paratypes are all immature, the largest specimen meas-
uring 10.3 mm in height. No additional material is known.
Discussion: Calliostoma sanjaimense is most closely
related to C’. iridium but has a sturdier shell, has strongly
beaded cords on the early whorls, which are lacking in
C. iridium, and has fewer and more prominent basal
cords. The immature paratype specimens show brilliant
metallic iridescence of green and yellow on the smooth
narrow interspaces between the spiral cords of the early
whorls, but this luster has evidently faded in the holotype.
Calliostoma veleroae McLEan, spec. nov.
(Plate 62, Figure 12)
Diagnosis: Whorls concave, spiral cording consisting of
a projecting peripheral cord and 6 evenly beaded cords
on the last whorl; base imperforate, but with a shallow
depression, base with about 18 low cords, beaded near the
columella. Color yellowish with brown flammules, basal
Page 424
cords with alternating light and dark markings. Height,
15.9, diameter, 17.0 mm (holotype).
Description: Shell of moderate size, sturdy, sides of
whorls concave, imparting a concave slope to the sides of
the shell. Postnuclear whorls 8, periphery sharply angu-
late, projecting, actually composed of 3 finely beaded
cords, 2 at the edge of the periphery and 1 just above.
Strong subsutural cord lacking. There are 3 beaded inter-
mediate cords on the 3 whorl, increasing by the addition
of intercalary threads that gradually increase in size and
become beaded until there are 6 cords and an equal
number of intercalary threads on the final whorl. Base
slightly convex, imperforate but with a hollow excavation
near the columella; base with 18 low spiral cords, the
interspaces of nearly equal width; innermost cords faint-
ly beaded in a radial pattern of growth. Color yellowish
with brown flammules, basal cords with alternating light
and dark markings.
Type Material: Holotype, LACM-AHF 1271.
Type Locality: Three miles south of Isla Ladrones, Pana-
ma, 7°49’ N, 82°23’30” W, 54 fms., Velero III station
943-39, 27 March 1939. The holotype, the only known
specimen, has a broken lip and a hole in the base of the
shell; the color is evidently somewhat faded.
Discussion: Calliostoma veleroae stands alone among
west American species in having a concave outline and a
sharply carinate periphery. It also differs from most of the
species discussed here in lacking a relatively strong sub-
THE VELIGER
Vol. 12; No. 4
sutural cord. A species of somewhat similar proportions
in the Caribbean fauna is Calliostoma aurora Daut, 1888,
but that species has a smooth, nearly concave base.
Calliostoma veleroae is named in honor of the Velero
III, the vessel of the late Captain G. Allan Hancock,
whose collecting expeditions formed the basis of the rich
molluscan material in the Allan Hancock Foundation
collection.
Calliostoma keenae McLean, spec. nov.
(Plate 62, Figures 13 to 16)
Diagnosis: Whorls and periphery rounded, spiral cording
unbeaded until the 4" whorl, final whorl with numerous
raised, finely beaded spiral cords, interspaces of equal
width; base imperforate, with about 12 low, weakly
beaded cords. Color, drab green or yellow brown with
brown flammules. Height, 14.9, diameter, 15.5 mm (holo-
type).
Description of Holotype: Shell of medium size, whorls
markedly convex, base of shell delimited by a thicker
spiral rib, postnuclear whorls 7. Sculpture of first to 3"
postnuclear whorls consisting of 3 raised cords, a trace of
beading appearing on the uppermost cord on the 4%
whorl; the middle of the 3 original cords persists as a
slightly more prominent cord on all succeeding whorls.
Additional cords arise from intercalary threads until, on
the final whorl, there are 15 cords between the basal
cord and the suture; interspaces are about as wide as
the cords and the uppermost cords are the more strongly
Explanation of Plate 62
Figure 1: Calliostoma iridium Dat, 1896. Holotype, USNM
122957. Gulf of Panama. Height, 21.5, diameter, 19.2mm. X 1.5
Figure 2: Calliostoma nepheloide Dati, 1913. Holotype, USNM
96637. Panama Bay. Height, 25, diameter, 22 mm. X 1.5
Figures 3 and 4: Calliostoma nepheloide. SODNHM 39799. Man-
zanillo, Mexico. Height, 20, diameter, 20 mm. XxX 1.5
Figure 5: Calliostoma nepheloide. AHF 618-37. San Jaime Bank,
Mexico. Height, 11.7, diameter, 11.6 mm. xX 15
Figure 6: Calliostoma fonki (Putrpri1, 1860). AHF 802-38.
Charles Island, Galapagos. Height, 17.5, diameter, 16.2mm X 2
Figure 7: Calliostoma fonkii. AHF 371-35. Independencia Bay,
Peru. Height, 9.1, diameter, 8.0 mm. X 4
Figures 8 and g: Calliostoma gordanum McLean, spec. nov. Holo-
type, CAS 13271. Gorda Banks, Mexico. Height, 19.6, diameter,
20.0 mm. XK 15
Figure 10: Calliostoma gordanum. AHF 531-36. San Francisquito
Bay, Mexico. Height, 13.2, diameter, 13.3 mm. X 1.5
Figure 11: Calliostoma sanjaimense McLEan, spec. nov. Holotype,
LACM-AHF 1269. San Jaime Bank, Mexico. Height, 20.0, dia-
meter, 18.4 mm. X 2
Figure 12: Calliostoma veleroae McLEan, spec. nov. Holotype,
LACM-AHF1271. Isla Ladrones, Panama. Height, 15.9, diameter,
17.0 mm. X 2
Figures 13 and 14: Calliostoma keenae McLean, spec. nov. Holo-
type, LACM-AHF 1272. Off Laguna Beach, California. Height,
14.9, diameter. 15.5 mm. X 3
Figure 15: Calliostoma keenae. AHF 618-37. San Jaime Bank,
Mexico. Height, 8.7, diameter, 8.9 mm. X 3
Figure 16: Calliostoma keenae. AHF 921-39. Clarion Island, Mexi-
co. Height, 10.5, diameter, 10.1 mm. X 3
Figure 17: Calliostoma jacquelinae McLean, spec. nov. Holotype,
SDNHM 51299. Santa Cruz Island, Galapagos. Height, 11.3, dia-
meter, 10.0 mm. XK 3
Figures 18 and 19: Calliostoma santacruzanum McL&an, spec. nov.
Holotype, SDNHM 51301. Santa Cruz Island, Galapagos. Height,
7.0, diameter, 6.9 mm. xX 5
Tue VELIcER, Vol. 12, No. 4 [McLean] Plate 62
Figure 6 Mgue Ue
Figure 14
Figure 16 Figure 18 i
Vol. 12; No. 4
THE VELIGER
Page 425
beaded. Base imperforate, basal cords about 12, inter-
spaces slightly wider than cords, particularly near the
columella where the cords are broader and more widely
spaced. Color, yellow brown with slightly darker flam-
mules, more pronounced darker and lighter areas at the
basal cord, base uniform yellow brown.
Type Material: Holotype, LACM-AHF 1272.
Type Locality: 58 Fathom Bank. 12 miles off Laguna
Beach, Orange County, California, 33°23’47” N, 117°
59’47” W, 58-60 fms., Velero IV station 1680-49, 12
February 1949.
Referred Material: Seven lots as follows: AHF 1254-41,
8 miles SW of Cedros Island, Baja California, 65 fms.;
AHF 1253-41, 8 miles W of Cedros Island, 65 fms.; AHF
1261-41, 4 miles N of Dewey Channel (Cedros Island),
24 - 25 fms.; AHF 618-37, San Jaime Bank, off Cape San
Lucas, Baja California (Plate 62, Figure 15); LACM A375,
Clarion Island, Revillagigedo Islands, Mexico, 30 fms.;
AHF 918-39, Sulphur Bay, Clarion Island, 45 - 60 fms.;
AHF 921-39, N of Clarion Island, 35-56 fms. (Plate
62, Figure 16). Two specimens are in the lot from San
Jaime Bank, but the other lots consist of a single speci-
men each. The holotype is the largest specimen. Only the
holotype and the 2 specimens from San Jaime Bank were
live-collected. Calliostoma keenae ranges from Laguna
Beach, California (33°12’N) to the Revillagigedo Is-
lands, Mexico (18°18’ N). It is chiefly a species of the
Californian province since its distribution encompasses
the entire outer coast of Baja California.
Discussion: Calliostoma keenae differs chiefly from east-
ern Pacific Calliostoma species of similar proportion in
having strong spiral cords on the early whorls that are
nearly devoid of beading. Calliostoma supragranosum
CarPENTER, 1864, a shallow water species of the Califor-
nia province, differs in having a greater number of ribs
that are strongly beaded on the early whorls. Calliostoma
leanum (C.B. Apams, 1852) has thicker, more strongly
beaded spiral cording. Calliostoma turbinum Datu, 1896,
an offshore species of southern California, has fewer
spiral cords and has a brassy metallic luster not shown
in C. keenae.
Calliostoma keenae is dedicated to Dr. Myra Keen of
Stanford University, whose warm and friendly manner
has been an inspiration to all workers in malacology.
Calliostoma jacquelinae McLEan, spec. nov.
(Plate 62, Figure 17)
Diagnosis: Whorls flat sided, spiral cording consisting
of 2 strong peripheral cords, a strong subsutural cord and
about 8 intermediate cords of varying strength, base im-
perforate, with numerous fine cords, the 4 cords near the
columella more prominent. Color light tan, all surfaces
highly opalescent with lavender and green. Height, 11.3,
diameter, 10.0 mm (holotype).
Description of Holotype: Shell relatively small for the
genus, thin, whorls flat sided, showing numerous growth
scars. Postnuclear whorls 7; periphery defined by 2
strongly projecting, heavily beaded cords with a thin
intercalary thread between; immediate subsutural cord
stronger than other cords of the body whorl; on the 2"4
and 3 postnuclear whorls there are 3 spiral cords and
faint axial sculpture producing a cancellate pattern, the
axial cords become obsolete by the 4 whorl and inter-
calary cords arise so that there is a total of 7 cords
between the subsutural cord and the 1* peripheral cord;
these cords are of varying strength and spacing, some are
weakly beaded and others are nearly smooth. On the
imperforate base there are 4 prominent, slightly beaded
cords near the columella and about 15 fine irregular
unbeaded cords between these 4 cords and the lower
peripheral cord defining the base.
Type Material: Holotype, SDNHM 51299, 1 paratype,
SDNHM 51300, 1 paratype, LACM 1274, 1 paratype,
USNM 679553.
Type Locality: South Academy Bay, Santa Cruz Island,
Galapagos Islands, Ecuador, 0°45’S, 90°20’ W, 150m
[82 fms.], dredged by André and Jacqueline DeRoy, 39
May and 10 June, 1969.
Discussion: Calliostoma jacquelinae with its opalescent
surface devoid of color pattern and its 2 strong peri-
pheral cords is distinct from all other eastern Pacific
species. Calliostoma schroedert CLENCH & Acuayo, 1938,
in the Caribbean province, resembles this species but
lacks sculpture other than the 2 peripheral cords.
The species is dedicated to Mrs. Jacqueline DeRoy
whose collecting has done much to increase our knowledge
of Galapagos Islands mollusks.
Page 426
Calliostoma santacruzanum McLEan, spec. nov.
(Plate 62, Figures 18, 19)
Diagnosis: Whorls flat sided, final whorl! slightly convex,
base narrowly umbilicate, spiral cording consisting of 8
beaded intermediate cords, a strong peripheral cord and
numerous unbeaded basal cords. Height, 7.0, diameter
6.9 mm (holotype).
Description of Holotype: Shell relatively small, post-
nuclear whorls 7, base narrowly umbilicate, base defined
by a single peripheral cord. On the first 3 postnuclear
whorls there are 3 strongly beaded spiral cords. Inter-
calary cords arise between these cords resuiting in 8
beaded cords of slightly variable size on the final whorl;
the immediate subsutural cord is slightly more prominent.
On the base only the cord bordering the umbilicus is
beaded, adjacent to it are 3 broad cords, followed by 12
narrow cords, separated by incised grooves. Color light tan
with broad brownish maculations, strongly marked along
the peripheral cord, green and yellow iridescence showing
on the smooth areas between the cords on the early whorls.
Whorls flat sided, although a slight concavity is evident
at the third whorl and the final whorl shows a slight
convexity.
Type Material: Holotype, SDNHM 51301.
Type Locality: South Academy Bay, Santa Cruz Is-
land, Galapagos Islands, Ecuador, 0°45’S, 90°20’ W,
25 fms., dredged by André and Jacqueline DeRoy, 10
June 1968. Efforts to obtain additional material have so
far been unsuccessful.
Discussion: Although only one specimen is known,
which is probably immature, Calliostoma santacruzanum
is easily distinguished from other eastern Pacific species.
It is the only umbilicate species other than the low-spired,
broadly umbilicate C. rema Stronc, Hanna & HERTLEIN,
1933.
THE VELIGER
Vol. 12; No. 4
LITERATURE CITED
Ciencu, WiLuiaM James & RutH Drxon TURNER
1960. The genus Calliostoma in the Western Atlantic.
Johnsonia 4 (40) : 1 - 80; plts- 1 - 56 (25 November 1960)
Dai, WILLIAM HEALEY
1896. Diagnoses of new species of mollusks from the west coast
of America. Proc. U.S. Nat. Mus. 18: 7-20
(April 1896)
1902. _ Illustrations and descriptions of new, unfigured, or im-
perfectly known shells, chiefly American, in the U.S. National
Museum. Proc. U.S. Nat. Mus. 24: 499 - 566; plts. 27 - 40
(March 1902)
1908. Reports on the dredging operations off the west coast
of Central America to the Galapagos, to the west coast of
Mexico, and in the Gulf of California. . XIV. The
Mollusca and Brachiopoda. Bull. Mus. Comp. Zool.,
Harvard 43 (6): 205 - 487; 22 plts. (October 1908)
1909. | Report on a collection of shells from Peru, with a sum-
mary of the littoral marine Mollusca of the Peruvian zoological
province. Proc. U.S. Nat.Mus. 37 (1704): 147 - 294; plts.
20 - 28 (24 November 1909)
1925. Illustrations of unfigured types of shells in the collection
of the United States National Museum. Proc. U.S. Nat.
Mus. 66 (2554) : 1-41; plts. 1 - 36 (22 September 1925)
Keen, A. Myra
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i-xi + 624 pp.; illus.
Stanford, Calif. (Stanford Univ. Press)
ParKER, Rospert H.
1964. | Zoogeography and ecology of some macro-invertebrates,
particularly molluscs, in the Gulf of California and the conti-
nental slope off Mexico. Vidensk. Medd. Dansk naturh.
Foren. 126: 1 - 178; 15 plts.; 29 text figs.; 7 tables (17 Febr. ’64)
Puteri, RupoLF AMANDUS
1860. Reise durch die Wiiste Atacama auf Befehl der chileni-
schen Regierung im Sommer 1853-54. Halle, Eduard An-
ton; 192+62 pp.; 27 plts.
Pitspry, HENRY AuGuUSTUS
1888-1898. Manual of Conchology. Philadelphia, 10 - 17
Vol. 12; No. 4
THE VELIGER
Page 427
An Annotated Bibliography of References to Marine Mollusca
of the Northern State of Sonora, Mexico °
BY
CAROL SKOGLUND
Department of Zoology, Arizona State University, Tempe, Arizona 85281
THE FOLLOWING LIST covers references to Mollusca along
the coast of the Mexican State of Sonora from Cape
Tepoca (Puerto Lobos) to the upper reaches of tidal
water in the estuary of the Colorado River. Six separate
places have been visited along this coastline since 1869
(see Table 1).
Dr. Edward Palmer, a botanist, was the first collector
in the area. In 1869 he traveled by boat from Yuma to
Guaymas, Mexico, collecting as he went (McVaucu,
1956). The first malacologist was H. N. Lowe (1934,
1935) who collected extensively in the Puerto Penasco
area in 1934. Starting in 1948, Dr. S. S. Berry (1953)
made several trips to Puerto Penasco and Choya Bay.
Both men described new species as a result of their find-
ings. Dr. P. Pickens collected opisthobranchs intensively in
the same area from 1963 to 1966. His work resulted in
descriptions of 19 new taxa by Marcus (1967).
The Hancock and Albatross Expeditions were the only
ones by sea to reach the eastern shore of the head of the
Gulf. Some of the material gathered by dredging was later
included in articles by specialists in certain families (Rost,
1955))e
Since most references have been to Cholla Bay, this
was used. Government maps now show this as Choya
Bay, which apparently was the original name. Other
names of places have been updated in so far as possible.
Where doubt of actual location exists, the original text is
given.
The references have been grouped as follows:
Descriptions of new taxa
Range extensions
Lists of species
Locality descriptions
General references
Ancillary references
Later citations of these same works have been omitted
from the list.
The author wishes to thank Dr. Dwight W. Taylor, for-
merly of the Department of Zoology, Arizona State Uni-
versity (and now at the San Diego Natural History
Collectors and Expeditions with Areas Visited
m Collector
ES
1869 Palmer
1889 Albatross
1890 Orcutt
1897! Boundary Comm.
1905 Rhoads
1907 Sykes
1909 Lumholtz
1915 Murphy
1931 Ives
1934 Lowe
1937 Huffman
Hancock Exped.
1940 Hancock Exped.
1941 Stanford Exped.
1944 Gifford
1945 Gifford
1947 Baker
1948 Berry
1949 Berry
1950! Bessom
1951 Thurlow
1952 Berry
1954 Mousley
Emerson
Turver
1955 French
Chace
1956 Rogers
Wright
Berry
Moore
1957 Chace
DuShane
Shasky
Colorado river
Table 1
XOX ew
xX X
mouth area
El Golfo de
Santa Clara
Adair Bay
x
Cholla Bay
x X X
xX XXX KK XK XK KK XK XK XK XK XK XK XK XK OX OX
Puerto Penasco
xX XK KK XK XK KK XK XK
x
(Puerto Lobos)
Cape Tepoca
x
Page 428
Colorado river
mouth area
El Golfo de
Santa Clara
Puerto Penasco
Cape Tepoca
(Puerto Lobos)
Adair Bay
q Collector
P
1958 Chace
1959 Chace
1959: Hall
1963 Pickens
1964 Brown x
1964 Donohue
Hanselman
Johnson, M.
Pickens
Paine
Rice
1965 Pickens
1966 Pickens
Farmer
Skoglund x
1967 DuShane x
1968 Johnson, W. S.
X X | Cholla Bay
x X
xX XX
XXxXXXXXXXK &
x
' publication date
Museum, San Diego, California) for many suggestions
and for reading the manuscript.
DESCRIPTIONS or NEW TAXA
Berry, SAMUEL STILLMAN
1945. Two new chitons from the Gulf of California. Amer.
Midld. Nat. 34 (2): 491 - 495; 18 figs. (23 August 1945)
Chaetopleura (Pallochiton) euryplax, new species, taken under
stones at Adair Bay by E. C. Huffman in 1937.
A new West Mexican prosobranch mollusk parasitic on
Amer. Midld. Nat. 56 (2): 355-357; 2 figs.
(October 1956)
Turveria encopendema, new genus and species, taken on Encope
grandis on outer strand of Cholla Bay by Mr. and Mrs. Harry R.
Turver, 1 May 1954, and by S.S. Berry in March 1949 and on
May 13, 1952.
1956.
echinoids.
1956. Diagnoses of new eastern Pacific chitons. Leafl. in
Malacol. 1 (13): 71-74 (19 July 1956)
Nuttallina crossota, Lepidozona subtilis, new species, and Stenoplax
(Maugerella) conspicua sonorana, new subspecies, taken at Norse
Beach near Cholla Bay by S.S. Berry in March 1948 and March
1949.
1957. Notices of new eastern Pacific Mollusca. I. Leafl. in
Malacol. 1 (14): 75 - 82 (19 July 1957)
Lithophaga (Labis) attenuata rogersi, new subspecies, taken by Mark
Rogers and Fred Wright in December 1956 at Cholla Bay, and
THE VELIGER
Vol. 12; No. 4
Acmaea stanfordiana, new species, taken at nearby Pelican Point by
S.S. Berry in March 1948.
1958. Notices of new Eastern Pacific Mollusca. — II.
Leaflets in Malacol. 1 (15): 83-90 (28 March 1958)
Tiphyocerma preposterum, new genus and species, taken by S.S.
Berry at Norse Beach in May 1952, and Olivella fletcherae, new spe-
cies, taken in March 1948,
1960. Notices of new eastern Pacific Mollusca — IV.
Leafl. in Malacol. 1 (19): 115 - 122 (31 December 1960)
Acmaea acutapex, new species, taken at “Punta Cholla, W. of
Puerto Pefiasco ...” by S.S. Berry in March 1948; Neosimnia vid-
leri tyrianthina, new subspecies, Berry and Moore, Cholla Bay, 1956;
Mitra (Tiara) lindsayi, new species, off Puerto Pefasco, 10 fms., H.
N. Lowe, February 1934.
1964. Notices of new eastern Pacific Mollusca. — VI.
Leafi. Malacol. 1 (24): 147 - 154 (29 July 1964)
Melampus mousleyi, new species, found in the upper estero of Cholla
Bay by S.S. Berry in March 1948.
CamMPBELL, G. Bruce
1964. New terebrid species from the eastern Pacific (Mollusca:
Gastropoda). The Veliger 6 (3): 132 - 138; plt. 17
(1 January 1964)
Terebra (Strioterebrum) adairensis, new species, found at Adair Bay
by Dorothy Brown.
Dati, WituiamM HEaLey
1891. Scientific results of explorations by the U.S. Fish Com-
mission steamer Albatross. XX. -— On some new or interesting
West American shells obtained from the dredgings of the U. S.
Fish Commission steamer Albatross in 1888 and from other
sources. Proc. U.S. Nat. Mus. 14 (849): 173-191; plts.
5-7 (24 July 1891)
Eupleura var. limata, described from the head of the Gulf of Cali-
fornia near the estuary of the Colorado River.
1894. On some species of Mulinia from the Pacific Coast.
The Nautilus 8 (1): 5-6; plt. 1 (2 May 1894)
Mulinia coloradoensis, new species, head of the Gulf of California
in the estuary of the Colorado River, found by Dr. E. Palmer.
1910. New shells from the Gulf of California. The Nauti-
lus 24 (3): 32 - 34 (2 August 1910)
Cymatium adairense, new species, found off Adair Bay.
1913.
Proc. U.S. Nat. Mus. 45 (2002) : 587 - 597
Amphissa (Cosmioconcha) palmeri and Erycina colpoica, new spe-
cies, from the head of the Gulf of California, found by Dr. E.
Palmer.
Diagnoses of new shells from the Pacific Ocean.
(11 June 1913)
Notes on the Semelidae of the West Coast of America,
including some new species. Proc. Acad. Nat. Sci. Phila-
delphia 67; 25 - 28 (2 March 1915)
Abra tepocana, new species, taken “off Cape Tepoca, Lower Cali-
fornia.”
1915.
Vol. 12; No. 4
THE VELIGER
Page 429
DuSuaneg, HELEN & James Hamitton McLEAN
1968. Three new epitoniid gastropods from the Panamic prov-
ince. Los Angeles County Mus. Contr. Sci. 145: 6 pp.; 6
figs. (14 June 1968)
Epitonium (Asperiscala) huffmani, new species, found at Cholla Bay
by Al Huffman in April 1937.
Lowe, HERBERT NELSON
1935. | New marine Mollusca from West Mexico, together with
a list of shells collected at Punta Penasco, Sonora, Mexico.
Trans. San Diego Soc. Nat. Hist. 8 (6): 15 - 34; plts. 1-4
(21 March 1935)
A list of 299 species, including 3 new species: Leda (Adrana) penas-
coensis, Turbonilla (Ptycheulimella) penascoensis, and Clathrodrillia
pilsbryi, which were dredged at 10 fms. by Lowe in 1934.
Marcus, EvELINE DU Bors-REYMOND & ErRNsT Marcus
1967. American opisthobranch mollusks. Studies in tropical
oceanography (Univ. Miami Inst. Marine Sci., Miami, Flori-
da), no. 6, 256 pp.
An excellent account of material collected by Peter Pickens and his
associates from November 1963 to April 1966. This work includes
descriptions of 2 new genera and species, 14 new species, and 3 new
subspecies. Thirty-nine species are reported from Puerto Pefiasco,
5 from Cholla Bay, and 9 from Puerto Lobos.
Pitspry, Henry AuGcusTus
1910. A new species of Marinula from the head of the Gulf of
California. Proc. Acad. Nat. Sci. Philadelphia 62: 148 - 149;
1 fig. (23 May 1910)
Marinula rhoadsi, new species, from “Hardie River, Lower Califor-
nia,” found by S. N. Rhodes in February 1905.
Pitspry, Henry Aucustus « AxEL ADoLF OLssoNn
1945. _ Vitrinellidae and similar gastropods of the Panamic prov-
ince. Part I. Proc. Acad. Nat. Sci. Philadelphia 97: 249 to
277; pits. 22 - 30 (27 December 1945)
Teinostoma politum ultimum, new subspecies, from ‘Rocky Bluff,
Bahia de Adair, Sonora ...,” collected by Earl C. Huffman.
SHasky, Donatp R.
1961. | New deep water mollusks from the Gulf of California.
The Veliger 4 (1): 18-21; plt. 4, figs. 1-10 (1 July 1961)
Trigonostoma campbelli, new species, from off Puerto Penasco in 10
fms., collected by H.N. Lowe in February 1934.
STEARNS, ROBERT EDWARDS CARTER
1893. Description of a new species of Nassa from the Gulf of
California. The Nautilus 7 (1): 10-11 (7 June 1893)
Nassa brunneostoma, new species, from the Gulf of California near
the mouth of the Colorado River, collected by Dr. E. Palmer.
RANGE EXTENSIONS
Cate, Crawrorp NEILL
1969. A revision of the eastern Pacific Ovulidae.
ger 12 (1): 95-102; plts. 7- 10; 3 maps
The Veli-
(1 July 1969)
The type locality for Jenneria pustulata (Licutroot, 1786) is
designated as Cholla Bay, Punta Pefiasco, Sonora, Mexico. Florence
Thurlow found the species at Cholla Bay in sand in April 1951
(personal communication). Simnia rufa (Sowersy, 1832) and S.
aequalis aequalis (SowERBY, 1832) are reported from Adair Bay.
Neosimnia quaylei Lowe, 1935 and N. vidleri tyrianthina Berry,
1960, become synonyms of S. a. aequalis.
1969. The eastern Pacific cowries. The Veliger 12 (1):
103 - 119; plts. 11-15; 3 maps (1 July 1969)
Erosaria albuginosa (Gray, 1825) is reported from Puerto Pefiasco.
Dai, WiLutiAM HEaLey
1897. | Report on the mollusks collected by the international
boundary commission of the United States and Mexico, 1892-
1894. Proc. U.S. Nat. Mus. 19 (1111): 333-379; plts.
31 - 33 (27 January 1897)
Tivela crassatelloides found near the mouth of the Colorado River
at the mouth of Rio Hardy.
DALL, WILLIAM HEALEY, & PAUL BARTSCH
1909. A monograph of the West American pyramidellid mol-
lusks. U.S. Nat. Mus. Bull. 68: 1 - 258; 30 plts.
(13 December 1909)
Pyramidella mazatlanica dredged off Cape Tepoca at the U.S.
Bureau of Fisheries station 3019 in March 1889 by the Albatross
(Townsend, 1901).
DonoHUuE, JERRY
1966. The range of Trivia myrae CAMPBELL. The Veliger
9 (1): 35-36; 1 map (1 July 1966)
Found at Puerto Pefiasco by Nora Donohue in April 1964.
FarMER, WESLEY MERRILL
1967. Notes on the Opisthobranchia of Baja California, Mexi-
co, with range extensions - II. The Veliger 9 (3) : 340 - 342;
1 text fig. (1 January 1967)
Chromodoris banksi and Dirona picta were found one mile south
of Puerto Pefiasco in February 1966.
Husss, Cary Leavitt « RoBERT RusH MILLER
1948. The Great Basin with emphasis on glacial and _post-
glacial times. IT. The zoological evidence. Bull. Univ. Utah
38 (20) : 17 - 166; 1 map (30 June 1948)
Cerithidea albonodosa found above the present tide level near the
lower Rio Hardy (p. 111).
McLean, James HamMILTON
1967. | West American species of Lucapinella. The Veliger
9 (3): 349-352; plt. 49; 3 text figs. (1 January 1967)
Lucapinella milleri reported from Puerto Penasco. A. Huffman.
Murpny, Ropert CUSHMAN
1917. Natural history observations from the Mexican portion
of the Colorado Desert. Proc. Linn. Soc. New York (28-29):
43 - 114 (11 December 1917)
Cerithidea sacrata Goutp found in the area of Laguna Salada, Baja
California in March 1915 by Murphy.
Page 430
Suasky, Donatp R.
1959. Range extensions for gastropods of tropical West Amer-
ica. Minutes Conch. Club South. Calif. (186): 15-18
Mitrella millepunctata range extension to Cholla Bay.
1961. Notes on rare and little known Panamic mollusks. The
Veliger 4 (1): 22-24; plt. 4, figs. 11-16 (1 July 1961)
Tenaturris nereis taken at Puerto Penasco by Shasky.
STEARNS, Ropert Epwarps CarTER
1894. The shells of the Tres Marias and other localities along
the shores of Lower California and the Gulf of California.
Proc. U.S. Nat. Mus. 17 (996): 139 - 204.
Cardium elatum found at Pinacate Bay [Adair Bay?] by Dr. E.
Palmer (p. 151).
Stronc, ArcuipaLp McCuiure « Hersert NeEtson Lowe
1936. | West American species of the genus Phos. Trans. San
Diego Soc. Nat. Hist. 8 (22): 305 - 320; plt. 22
Phos gaudens taken by H.N. Lowe in 10 fms. off Puerto Penasco.
LISTS or SPECIES
BErry, SAMUEL STILLMAN
1956. See locality descriptions
Burcu, Beatrice LaRue
1967. | Murex shells. Cholla Chatter (Cholla Bay Sports-
men’s Club, Phoenix, Arizona) 10 (5): 8-10 (May 1967)
A list of Muricidae to be found in the Cholla Bay area with infor-
mation on life cycles of some species.
Burcu, JoHN Quincy
1967. The olive shells of Cholla Bay. Cholla Chatter
(Cholla Bay Sportsmen’s Club, Phoenix, Arizona) 11 (10):
8-9; 1 plt. (November 1967)
A review of the seven members of the Olividae found at Cholla Bay.
DuSuHane, HELEN « ELLEN BRENNAN
1969. _A preliminary survey of mollusks for Consag Rock and
adjacent areas, Gulf of California, Mexico. The Veliger
11 (4): 351 - 363; 1 map (1 April 1969)
Seven species collected by DuShane at El Golfo de Santa Clara in
November 1967 are listed.
Girrorp, Epwarp WINsSLow
1945. Archaeology in the Punta Pefiasco region, Sonora.
Amer. Antiq. 11: 215-221; fig. 29 (May 1945)
A list of shells found at old camp sites and on nearby dunes.
HErTLEIN, LEo Georce & WILLIAM KerirH EMERSON
1956. Marine Pleistocene invertebrates from near Puerto Pen-
asco, Sonora, Mexico. Trans. San Diego Soc. Nat. Hist.
12 (8): 154-176; plt. 12; 2 maps (7 June 1956)
A list of fossil shells taken at Puerto Penasco and Pelican Point
(Cholla Bay) with comparisons to present day fauna.
THE VELIGER
Vol. 12; No. 4
Hornapay, WILLIAM T.
1909. Camp-fires on desert and lava.
Sons, New York, 366 pp.
Five species found on the shores of Adair Bay by Godfrey Sykes in
1907 (p. 240).
Charles Scribner’s
Jounson, MyrtLe EvANGELINE
1968. The cone shells of Cholla Bay. Cholla Chatter
(Cholla Bay Sportsmen’s Club, Phoenix, Arizona) 11 (12):
4-7; figs. 1-4 (December 1968)
Lists cones found in the Penasco-Cholla Bay area.
Keen, A. Myra
1947. Species of mollusks collected by the Stanford Expedition,
1941, to be added to the list published by H.N. Lowe, 1935
(Trans. San Diego Soc. Nat. Hist., vol. 8, no. 6, pp. 27 - 34).
Min. Conch. Club South. Calif. (75): 3-4 (Decemb. 1947)
[published anonymously but actually by A. Myra Keen as per
personal communication April 20, 1969.]
Sixty-one species to be added to the Puerto Pefiasco area list.
Lowe, H.N.
1934. See locality descriptions.
Marcus, E. « E. Marcus
1967. See descriptions of new taxa.
Rice, THomas C.
1964. Mexican shelling. Pacif. Northwest Shell News 4
(3, 4): 29-34 (Fall 1964)
A trip to Puerto Penasco and Cholla Bay in February 1964 with
George and Virginia Hanselman results in a preliminary list of 67
species.
1966. H & R expedition species check list. Pacif. North-
west Shell News 6 (1): 4, 13 - 22 (January 1966)
Lists 84 species from Puerto Penasco and 52 species from the Cholla
Bay area taken by Tom Rice in February 1964.
Suasky, Donatp R.
1958. See locality descriptions.
LOCALITY DESCRIPTIONS
Berry, SAMUEL STILLMAN
1956. _A tidal flat on the Vermilion Sea. Journ. Conchol.
24 (3): 81 - 84 (February 1956)
An excellent description of Cholla Bay; it includes the major species
to be found in each habitat from the back estuary to the low tide
line.
DuSHANE, HELEN
1957. Marine treasures from the beach at Punta Penasco.
Desert 20 (10): 17-20; 2 figs.; 2 maps (October 1957)
Collecting in Cholla Bay, with a photograph of 28 species found
by the DuShanes in April 1957 (personal communication) .
Vol. 12; No. 4
THE VELIGER
Page 431
Lowe, HERBERT NELSON
1934. On the Sonoran side of the Gulf. The Nautilus
48 (1): 1-4 (10 July 1934) ; 48 (2): 43-48 (15 Oct. 1934)
General account of a trip to Puerto Penasco in February 1934 with
information on habitats of mollusks seen.
Suasxy, DonaLp R.
1958. Collecting stations — Puerto Penasco, Son., Mexico.
Min. Conch. Club South. Calif. (176): 6 - 8; 1 fig. (April 1958)
Information on types of habitats and species to be found, from a
trip in March 1957.
GENERAL REFERENCES
FaRMER, WESLEY MERRILL
1968. _Tidepool animals from the Gulf of California. Wes-
word Co., San Diego, Calif.; 69 pp.; 170 figs.; 23 plts.
Drawings of many common mollusks of both El Golfo de Santa
Clara and Puerto Penasco.
Keen, A. Myra
1958. Sea shells of tropical West America; marine mollusks
from Lower California to Colombia. i- xi + 624 pp.; illus.
Stanford Univ. Press, Stanford, Calif. (5 December 1958)
A comprehensive work with references to ranges of many species to
the head of the Gulf of California and to Puerto Penasco.
ANCILLARY REFERENCES
Baker, Epwarp P
1947. The Bakers take a trip to Mexico. Min. Conch. Club
South. Calif. 72: 14-19 (August 1947)
An account of a shell collecting trip to Puerto Penasco and Cholla
Bay in January 1947
Berry, SAMUEL STILLMAN
1953. Molluscan notes from the Puerto Pefiasco region, Sonora
(abstract). Min. Conch. Club South. Calif. 131: 3 (Sept.)
Trips to the area by S.S. Berry in 1948, 1949, and 1952 resulted in
704 species being taken, many of which were undescribed.
1963. Diagnoses of new eastern Pacific chitons. II. Leafl.
in Malacol. 1 (22): 135-138 (29 March 1963)
Lepidozona pella, new species, from San Felipe, Baja California, is
compared with L. subtilis Berry, 1956 from Puerto Penasco.
Burcu, JoHN Quincy (ed.)
1949. _ Extracts from the Minutes of Conch. Club South. Calif.
92: 7
H. Turver spoke of collecting 300 species at Cholla Bay. The short
account mentions some of the species, and states that sizes are larger
than in the Guaymas area.
1950. Extracts from the Minutes of Conch. Club South. Calif.
104: 4 (September 1950)
L. Bessom gave an account of a trip to Puerto Pefiasco and Cholla
Bay.
1953. Extracts from the Minutes of Conch. Club South. Calif.
125: 6 (January/February 1953)
S. S. Berry spoke on “Molluscan notes from Puerto Penasco region.”
He stated that he has material from the region from Lowe, Keen,
Turver, Mousley, and the Fletchers.
1954. Extracts from the Minutes of Conch. Club South. Calif.
138: 40 (May 1954)
Louis B. Mousley found a large group of Cancellaria acuminata
live in Adair Bay in January 1954.
1954. Extracts from the Minutes Conch. Club South. Calif.
139: 60 (June 1954)
The Harry Turvers report good collecting at Puerto Pefiasco.
1955. Extracts from the Minutes of Conch. Club South. Calif.
152: 12 (November 1955)
Mr. and Mrs. Mead French gave a report on a recent trip to
Cholla Bay.
1959. Extracts from the Minutes Conch. Club South. Calif.
190: 14 (July/August 1959)
S. Ralph Hall found 2 dead Spondylus shells at Cholla Bay.
Burcu, THomas Apams (ed.)
1969. Cone shells of Cholla Bay. Cholla Chatter (Cholla
Bay Sportsmen’s Club, Phoenix, Arizona) 12 (1) :9; 1 fig.
Conus regularis found by Carol Skoglund at Cholla Bay boat landing
in August 1966.
1969. | Wormfish search. Cholla Chatter (Cholla Bay
Sportsmen’s Club, Phoenix, Arizona) 12 (6): 24-25 (June)
Cyclinella sp. found in the Cholla Bay estuary in May 1969.
Cuace, EMERY Perkins & Erste Marcaret Herpst CHACE
1967. | Conchological reminiscences. Recollections of Emery P.
Chace and Elsie M. Chace with the help of our notebooks.
San Diego Soc. Nat. Hist. 38 pp.
Collecting trips to Cholla Bay were made by the Chaces in 1955,
1957, 1958, and 1959.
GaRDNER, ERLE STANLEY
1969. Clamming at El Gulfo. Desert 32 (7): 12-15,
33 (July 1969)
Clam digging for food; no species names given.
Ives, RonaLp LorENz
1951. High sea levels of the Sonoran shore. Amer. Journ.
Sci. 249: 215 - 223; plts. 1-2; 2 figs. (March 1951)
Deposits of Chione cancellata and Turritella above present shoreline
used as indicators of past changes.
1959. = Shell dunes of the Sonoran shore. Amer. Journ. Sci.
257: 449-457; plts. 1-2; 3 figs. (June 1959)
Chione cancellata a component of the shell dunes.
1963. The problem of the Sonoran littoral cultures. Kiva
28: 28 - 32; 1 fig.
Page 432
THE VELIGER
Vol. 12; No. 4
Turritella leucostoma and Chione cancellata used to identify old
shorelines.
Jounson, MyrtLe EvANGELINE
1964. | My hobby — shell collecting. Cholla Chatter
(Cholla Bay Sportsmen’s Club, Phoenix, Arizona) 8(3): 14
to 15 (March 1964)
Collecting at Cholla Bay.
JoHNson, WILLIAM Scott
1968. Ecology and seasonal fluctuations of an intertidal Sar-
gassum-Hyale-Barleeia association at Punto Penasco, Mexico
(abstract) . Southwest. and Rocky Mt. Div., Amer. Assoc.
Adv. Sci. 44t* ann. meet. 28 April - 1 May 1968
LuMHOLTz Cari
1912. New trails jn Mexico.
York
Dosinia ponderosa seen at la Salada, near the mouth of the Colo-
rado in 1909 by Lumholtz (p. 256).
Charles Scribner’s Sons, New
Rost, HELEN
MacGrniTiz, GeorcE Esper & Nettie MacGIniTIEz
1949. Natural history of marine animals.
McGraw-Hill, New York.
473 pp.; illus.
Lamellaria orbiculata seen on white encrusting sponge at Puerto
Penasco (p. 372).
Orcutt, CHARLES RUSSELL
1891. A visit to Lake Maquata. West. Amer. Scient.
7 (59) : 158 - 164 (April 1891)
Cerithidea and clams of the genus Venus were found to the west-
ward of Lake Maquata.
PaInE, Rosert T.
1966. Function of labial spines, composition of diet, and size of
certain marine gastropods. The Veliger 9(1): 17-24;
2 text figs. (1 July 1966)
Observations on Acanthina angelica made at Puerto Penasco in
March 1964.
1955. A report on the family Arcidae (Pelecypods) .
Allan Hancock Pacif. Exped. 20 (2) :
17 text figs.
177 - 249; plts. 11 - 16;
(10 November 1955)
Offshore work in 1937 and 1940 at Puerto Penasco and at Cape
Tepoca. Anadara multicostata and A. alternata reported from
Penasco, and A. alternata from Tepoca.
LITERATURE CITED
McVaucu, RocEers
1956. | Edward Palmer, plant explorer of the American West.
Univ. Oklahoma Press, Norman, Okla. 430 pp.
TowNnsEND, C. H. (compiler)
1901. Dredging and other records by the U.S. Fish Commis-
sion steamer Albatross with the bibliography relative to the
work of the vessel.
Extr. U.S. Fish. Commiss. Rep. of
1900; Washington, D.C., pp. 387 - 562; plts. 1-7
Vol. 12; No. 4
THE VELIGER
Page 433
The Systematics and Some Aspects of the Ecology
of the Genus Dendronotus
(Gastropoda : Nudibranchia )
BY
GORDON A. ROBILLIARD
Department of Zoology and Friday Harbor Laboratories, University of Washington, Seattle, Washington 98105
\
(Plates 63 and 64; 28 Text figures)
INTRODUCTION
Tue cGENuS Dendronotus has had a long and chaotic
literary history, beginning in 1774 when Ascanius de-
scribed the type species under the name of Amphitrite
frondosa. MULLER (1776) incorporated this species into
his rather all-inclusive genus Doris as Doris arborescens.
Cuvier (1797) established the genus Tritonia and he
included A. frondosa as Tritonia arborescens in 1817. AL-
DER & Hancock (1845), on morphological bases, estab-
lished the genus Dendronotus as separate from Tritonia
with Doris arborescens Murr, 1776 as the type species.
Subsequently, the genus has been referred to as Dendro-
notus with the name of the type species D. frondosus
(Ascanius, 1774) being used for the first time in the
1904 edition of the Plymouth Marine Invertebrate Fauna
(Eutot, 1910).
During the late 18", 19 and early 20" centuries, the
species of Dendronotus, especially D. frondosus, were, with
only a few exceptions (ALDER & Hancock, 1845; VeErR-
RILL, 1870; MAcFarLanp, 1966), based on sketchy, in-
adequate descriptions and diagrams of one or a few
animals. Consequently, there are no reports on the range
of intraspecific variation of the various taxonomically
important characters, the radula excluded, for any species.
OpHNER (1934), in an effort to point out some of the
pitfalls facing opisthobranch taxonomists and what they
might do to avoid the same, states:
“... I think it is necessary to give good figures of the whole
animal, and not of details only ... as external shape and
features provide not only an immediate means of recognition,
but often also important systematic characteristics .... The
literature on nudibranchs is full of detailed descriptions but too
little of comparisons, which, however, are specially desirable
in this group, because of the difficulty of finding representative
characters in these soft-bodied animals ...”
During a comparative study of the nudibranch genus
Dendronotus from the San Juan-Puget Sound area, Wash-
ington, I found that there were apparently 3 more species
than were presently accepted names. In this paper I have
redescribed the 6 known and accepted species {D. fron-
dosus (AscaAntus, 1774), D. iris Cooper, 1863, D. robus-
tus VERRILL, 1870, D. gracilis Bapa, 1949, D. albus Mac-
FaRLAND, 1966, and D. subramosus MacFaranp, 1966}.
I have reinstated as well as redescribed two species, D.
dali Bercy, 1879 and D. rufus O’DonocHuE, 1921,
synonymized with D. frondosus by OpHNER (1936). I
consider D. venustus MacFarLanD, 1966 a synonym of D.
frondosus. Finally, I have described a new species of
Dendronotus.
For the sake of completeness, a diagnosis of the order,
family, and genus, drawn from the literature, is included.
I have included a definition or a figure or both of each
taxonomically significant character referred to in this
paper. Brief mention is made of a number of aspects of the
nudibranchs’ ecology and life history.
MATERIALS anp METHODS
Because of the distinct ecological differences of the local
species of Dendronotus, the animals were collected from
a number of different areas from August 1966 to June
1967 (see Appendix I for longitude and latitude of the
collection sites).
Dendronotus frondosus was collected from the moorage
floats in Friday Harbor and Mitchell Bay during August,
September, and early October of 1966 and again in April
and May of 1967. During late October and throughout
November, 1966, they were often found in the seawater
tables at the Friday Harbor Laboratories. They have been
Page 434
collected in February, 1967 and 1968, at Cantilever Pier
in 35 m of water.
From June, 1965 to May, 1967, Dendronotus iris was
collected incidentally while the research vessel, M. V.
“Hydah,” was otter trawling for flat fish in East Sound
and Bellingham Bay. Occasionally, it was dredged in
West Sound and Harney Channel. During June, 1967, I
collected 15 animals while SCUBA diving at Departure
Bay and Clarke Rock, Nanaimo, B. C.
Dendronotus subramosus was collected at Peavine Pass,
Edwards Reef, Lonesome Cove, and Low Island during
late August through to December, 1966 and in July, 1967.
Dendronotus rufus was obtained from the outlet of the
storm and sewer drain at Alki Point, Seattle during
November through February of 1966 to 1967 and 1967
to 1968, respectively. Dendronotus dalli was collected on
August 20, 1966; from October, 1966 to January, 1967,
and from January 1 to 4, 1968, on the Victoria Break-
water, Victoria. Dendonotus albus was collected at Lone-
some Cove, Brown Island, Collins Cove, and Edwards
Reef from September to December, 1966. The new species
of Dendronotus was obtained from Lonesome Cove, Brown
Island, the Cantilever Pier, and the Victoria Breakwater
from July, 1966 to January, 1967, although seldom in
abundance. All of the species were collected less commonly
in places other than those mentioned above.
The last 5 species I collected while SCUBA diving
because survival is about 5 per cent when dredged and
about 95% when collected while diving. Each animal
plus its substratum, usually a hydroid, was placed in a
plastic bag which could be tied shut and put in a canvas
collecting bag. Ecological data, such as location, depths,
currents, substratum, bottom type, possible food items, and
reproductive activity were recorded on a sheet of 1.5
mm white matte plastic with a soft lead pencil.
Each nudibranch was kept in an individual container
on the seawater table for about 2 days during which time
the fecal pellets were examined to determine what the
animal was eating prior to collection. A detailed descrip-
tion of each individual was then recorded on a prepared
worksheet, special attention being paid to body propor-
tions and dimensions, cerata, rhinophores, lip and veil
papillae, color, and the ecological data mentioned above.
A total of 15 Dendronotus rufus, 23 D. subramosus, 81
D. albus, 26 Dendronotus spec. nov., 28 D. iris, 32 D.
frondosus, and 53 D. dalh were thus described. Many
more specimens of each species were collected from May
to September, 1967 and from January to March, 1968.
They were checked to see if the above-mentioned charac-
ters fell within the range of variation described for the
species.
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Vol. 12; No. 4
To study the reproductive system, jaws and radula,
each animal was relaxed with succinylcholine (BEEMAN,
1968), killed with 5% seawater formalin, and stored in
5% seawater formalin with 0.25% Ionol C. P-40 (Rosi-
LIARD, 1969).
The ducts of the isolated genital system were separated
to facilitate study of the relative position and size of
component parts. The “female gland mass,” consisting
of the fertilization chamber, “albumin” gland, membrane
gland, and mucous gland (GuisE.in, 1965) was not drawn
because the component parts are very fragile and cannot
be easily separated. Furthermore, the size and shape of
this organ depend very much on the reproductive activity
of the genital system and are of minor taxonomic signifi-
cance.
The excised buccal mass, placed in a petri dish, was left
until all the tissue had decomposed. The radula was easily
extracted from jaws, cleaned, and mounted in a non-
resinous mounting medium (Turtox CMC-10). The jaws
were washed and preserved in 5% seawater formalin.
Both the radulae and the jaws were drawn using a
camera lucida and the drawings were supplemented by
reconstructions based on photographs.
Specimens of all the species studied have been depos-
ited in the California Academy of Sciences, the Smith-
sonian Institution, and the Canadian National Museum.
SPECIES
Introduction to Species Descriptions
I have tried to use large samples of each species, consisting
of animals from different localities, in order to account
for the considerable intraspecific variation that occurs in
the nudibranchs. Only living or well-relaxed, freshly pre-
served adult (or near-adult) animals have been described
because of the paucity of very young specimens. The
described animals are from a relatively limited area and
there may be even more variation throughout their whole
geographical range. ’
I have tried to compare and contrast each species with
the other species by describing quantitatively (or quali-
tatively, if more appropriate) the same structure in each
species. Where the description is quantitative, the first
number(s) represents the mode of the measurements or
counts, determined from all the animals used, followed by
the range of variation enclosed in parentheses (2 g., body
length, 5.0-6.0 (3.0-9.0) cm-L).
Within the description of each species, I have tried to
include the useful and accurate portions of earlier ac-
counts. Because earlier authors generally have not em-
Vol. 12; No. 4
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Page 435
\
ployed a consistent nomenclature when referring to the
anatomy and morphology of Dendronotus, I have applied
distinct terms to each taxonomically significant structure
and have defined and figured them. Although a few of
the terms are original, most have been used before; syno-
nyms and their references are included in Appendix II.
Except for the jaws, radula, and genital organs, the
internal organs are not described as they appear to be
very similar in all the species examined. For relatively
complete descriptions and accurate diagrams of the inter-
nal organs, the reader is referred to ALDER & HANCocK
(1845), Bercu (1894), OpHNnER (1936), and MacFar-
LAND (1966). The ganglia and nerves of the central ner-
vous system, although slightly different in each species,
are not described because they are difficult to isolate and
the differences do not appear to be significant on the
species level. BERcH (1879, 1894) and MacFarianp (op.
cit.) gave good descriptions and figures of the central
nervous system.
The choice of important taxonomic characters is neces-
sarily arbitrary (Mayr, 1965) but familiarity with a taxon
prepares one, potentially at least, to choose those charac-
teristics which will be most enlightening. After examining
7 species of Dendronotus, I have chosen several external
and internal characters, listed below, on which I have
based the species descriptions. These characters show the
greatest intraspecific consistency while being demonstrably
different interspecifically.
Six of the 9 named species of Dendronotus plus a new
species were found in the study area. The other species,
D. robustus and D. gracilis, are included for completeness,
the description being based on a compilation of the avail-
able literature. This latter type of description is relatively
unsatisfactory and should be used with some caution
because of differences of opinion between biologists con-
cerning the taxonomic significance of particular characters.
Synonymy and Taxonomic Remarks
Each synonymy cites only the first reference to a name or
new combination of names that were already in use. To
insure accuracy in the fairly exhaustive search for all the
names used for a species of Dendronotus, I have tried to
examine the original descriptions. When this has not been
feasible, all other references concerning the elusive de-
scription were utilized. Where it is appropriate, I have
made some remarks concerning the literary history of the
species in an effort to indicate why certain authors felt
new species should be established while others felt that
lumping of certain species would be more realistic.
Body Form, Dimensions, and Texture
The body of Dendronotus is limaciform or “sluglike,” but
differences in body proportions determine whether or not
the species is “delicate” or “heavy.” The length (L) of
the nudibranch is measured from the tip of the tail to
the anterior edge of the veil between the two medial veil
papillae when the animal is actively crawling (Figure 1).
CL 1.0cm
Figure 1
Ventral view of Dendronotus dalli, showing lip and veil papillae,
position of the mouth, and the foot
CE = cerata CL = clavus CP = crown papillae
= foot LI = lip papillae LP = lateral papilla
MGA = male genital aperture RST = rhinophore stalk
VP = veil papillae
The width (W) and height (H) are measured halfway
between the first and second pair of cerata over the cardiac
prominence, again when the animal is actively crawling
(Figures 2, 3). The height is the distance from the sub-
stratum to the top of the cardiac prominence. The width
is taken at the widest part of the body, exclusive of the
edge of the foot, at the same point. These dimensions
may vary more than 10% in a single animal because of
its plasticity.
Texture refers to the surface of the epidermis. In most
species, the epidermis is quite smooth, but in Dendronotus
frondosus, D. subramosus, and a few D. albus, there are
few to many conical (or subconical) papillae of various
sizes scattered over the dorsum, particularly the cardiac
prominence (Figure 3). They are usually capped with a
yellow to cream colored pigment.
In all the specimens examined, the anus and genital
orifice opened at approximately the same place. The
anus, often on a raised papilla, is located about halfway
along an imaginary line joining the bases of the 1* and
Page 436
\t g AP CE
Ay Z
VP SS yRst \ \ fe Ze
Fe
WibxS 2g : é.
S aL ES
(Fav Le Wsy NARS
—% Y V
ile a ANS
CP 1.0cm
Figure 2
Dorsal view of Dendronotus albus, showing the external features
of the body and appendages
AP = anal papilla CE = cerata
CP = crown papillae IL]? =
RSH = rhinophore sheath
VL = veil
CL = clavus
lateral papilla
RST = rhinophore stalk
VP = veil papillae
2-4 cerata on the right side. The genital openings are
located just anterior to the base of the 1* ceras on the
right side, usually from 4 to ? of the way up the side
(Figure 3).
3
‘\ i
ee
Wee
Gf
RW RST RSH
FGA =
MGA Z SS
s a CL
Zi aa
CP
1.0cm VP VL
Figure 3
Lateral view of Dendronotus subramosus, showing the genital
apertures and the cardiac prominence and papillae
AP = anal papilla BP = body papillae CE = cerata
CL = clavus CP = crown papillae
CPR = cardiac prominence - FGA = female genital aperture
MGA = male genital aperture RSH = rhinophore sheath
RST = rhinophore stalk VL = veil VP = veil papillae
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Vol. 12; No. 4
Foot
The foot (Figure 1) is briefly described, emphasis being
placed on the length, width, shape, and functional aspects.
The length (L) is the distance from the tip of the tail to
the most anterior edge of the foot, while the width (W)
is measured as the widest part of the foot. Both measure-
ments are taken while the animal is crawling. The discus-
sion of the functional aspects of the foot, particularly
in relation to the animal’s feeding habits, is based partly
on field and laboratory observations and partly on specu-
lation. Further studies are being made to determine the
relationship between the morphology of the foot and the
animal’s feeding habits.
Color
Color is one of the most variable characters in Dendro-
notus. However, if the variations are recognized and some
caution is exercised, color can be used as a guide in identi-
fying the species. In many cases, there are distinct patterns
(cf. D. subramosus) that occur consistently and allow one
to identify the animal even though the ground color is
different. As a final guide, color plates are presented
(Plates 63 and 64).
Cerata
The term cerata (Figures 2,3) should probably be reserved
for the dorsal appendages of the Aeolidiacea (Morton,
1958), but, because the term has been more or less ac-
cepted, and because it distinguishes between dorsal pro-
cesses (rhinophores and “cerata’”’), I have used it in this
paper. The possible function of the cerata as respiratory
organs is evident from the nomenclature that has been
applied by previous authors (see Appendix II) although
there has been no experimental work done to prove this.
Some aeolids have cerata with cnidosacs which presum-
ably are for protection (EpMuNps, 1966), but there
appear to be no cnidosacs in Dendronotus (PRuvot-Fot,
1954; THompson, 1960a). Other aeolids are able to
autotomize the cerata as a possible protective mechanism
(Epmunps, op. cit.; Garstanc, 1889; Stasek, 1967;
Tuompson, 1964), but the dendronotids appear unable
to accomplish this feat (unpublished observations). I have,
however, examined a number of specimens which have
lost parts of or whole cerata, rhinophores, or veil papillae,
probably from attacks by predators such as fish.
The branching patterns (Figure 4) and the size of the
cerata are relatively distinct and consistent within a spe-
cies. Each ceras has 1 - 5 main branches, defined as those
large branches arising directly from the body or from very
Vol. 12; No. 4
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Page 437
Figure 4
Branching patterns of cerata
A. “Fan-shaped” pattern with the left ceras showing all 3 branches
arising from a single stalk while the right ceras shows the lateral
branch arising first and the medial 2 arising above this.
B. “Rosette” pattern.
C. Primary (1°), secondary (2°), and tertiary (3°) branching of
a single ceras.
near the base of the ceras. From these main branches
arise a few to many secondary and tertiary branches
(Figure 4c). Normally the medial main branch is the
longest, the others becoming progressively shorter with
the lateral branch being the shortest. In some species, e. g.,
Dendronotus iris, it is difficult to distinguish the main
branches from large secondary branches and the choice
becomes arbitrary. There are two basic patterns which I
have called the “fan-shaped” pattern (Figure 4a) and
the “rosette” pattern (Figure 4b). The “fan-shaped”
ceras is oriented with the “fan” at right angles to the long
axis of the body. The “rosette” pattern consists of the
branches radiating from the top of a stout stalk. In a few
specimens, particularly of D. rufus, there are small tufts
called “accessory” cerata appearing in irregular groups
between the main cerata, especially over the cardiac
prominence.
The first pair of cerata usually is the largest with suc-
cessive posterior pairs becoming smaller and less branched.
The distance between successive pairs usually decreases
posteriorly. The reduction in the number of main branches
appears to be due to the loss of the lateral branches.
Rhinophore and Clavus
It has been generally thought that the rhinophores, par-
ticularly the clavus, “‘on the basis of their position, struc-
ture and innervation” (Koun, 1961, p. 300) are used as
chemoreceptors, especially for the detection of food. Most
of the experimental work with food extracts has failed to
show conclusively that the rhinophore is any more sensi-
tive to chemosensory stimulation than any other part of
the cephalic region (KoHn, 1961). However, WoLTER
(1967) has shown that the rhinophore is a chemosensory
structure, stimulated by food, in Archidoris tuberculata,
Polycera quadrilineata, Aeolidia papillosa, and Facelina
drummondt.
The 3 important components of the rhinophore are the
lateral papilla, the crown papillae, and the clavus (Fig-
ures 2, 3) (see Appendix II for synonyms). The presence
or absence, size, pattern of branching, and point of origin
on the rhinophore stalk are important in describing the
lateral papilla, the function of which is unknown. The
crown papillae, vertical prolongations of the rhinophore
sheath, vary in length, number, degree of branching and
arrangement on the sheath. The leaves of the clavus vary
in size, shape, number, and color. The overall shape of
the clavus is also important. In some cases, the position at
which the rhinophores are held while the animal is
crawling is distinctive.
Head, Lips, and Veil
The cephalic region of Dendronotus is very sensitive and
there is a strong indication that the veil papillae, and
possibly the lip papillae, serve as contact chemoreceptors
(unpublished observation) as do the oral tentacles of some
aeolids (Kon, 1961). As well as indicating the existence
of food, the veil papillae may warn the nudibranch of a
potential predator. When the animal crawls, the veil papil-
lae are held upward and forward. As soon as they con-
tact anything unfamiliar or different from the immediate
surroundings, the animal retracts and then cautiously
extends itself, exploring the area with the veil papillae.
If it is unpleasant, the nudibranch will turn away. If,
however, it is food, Dendronotus advances immediately
and begins to feed. In all the local species which feed on
hydroids, there are some indications that the lip papillae
Page 438
are used to orient the mouth and lips parallel to the long
axis of the hydroid stalk. The jaws may then be effectively
employed in holding the stalk while the radula rasps
through the perisarc, and the animal can withdraw the
coenosarc easily.
The most important taxonomic features of the head
region (Figures 1 to 3) are the number, degree of
branching, and arrangement of the veil and lip papillae.
The color of the lips is also distinctive in some cases. No
attempt was made to describe the inner labial ring and
other mouth armature described by MacFar.Lanp (1966).
The veil papillae have been arbitrarily defined as those
papillae which originate in a single row from the most
distal edge of the more or less distinct, horseshoe-shaped
veil. The lip papillae are all those papillae in the ventral
cephalic region not included in the above. In some species,
such as Dendronotus rufus, the lip papillae may be as
long as the shorter veil papillae and more branched, but
their position establishes their relationship. The lip pa-
pillae may be arranged in vaguely regular rows. It must
be emphasized that this definition of the papillae may be
artificial as it is based on position, not function.
To facilitate identification of any particular pair of veil
papillae they are numbered from medial to lateral, the
medial pair being pair no. 1 (Figures 2, 3).
Jaws
The jaws of Dendronotus, enclosing most of the buccal
mass, provide attachment for the jaw and radula muscles.
The functional portion of the jaw is the masticatory pro-
cess which is responsible for holding the prey and possibly
chops off pieces of the prey as it is pulled into the mouth
by the radula. Inferences about the species’ feeding
habits may be made from the structure of the jaw and its
size relative to the size of the animal.
The descriptions and diagrams (Figures 5, 7, 10, 13, 16,
19, 22, 25) attempt to point out differences in shape and
relative proportions of the jaw components. With the ex-
ception of Dendronotus iris, the jaws are fairly uniform
morphologically within the genus and one must exercise
caution in attempting to identify the species of Dendro-
notus from the jaws alone.
For a fairly complete description and excellent drawings
of the jaws of some species of Dendronotus, the reader is
referred to MacFarLanp (1966). He includes a detailed
description of the denticulation on the masticatory border
of the jaw, but I feel that this is not a particularly useful
taxonomic character, partly because of the difficulty in
examining the denticles, and partly because of the con-
siderable intraspecific variation and interspecific overlap
in number and shape.
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Vol. 12; No. 4
Figure 5
Measurements and components of a Dendronotus jaw
A. Inside lateral view B. Dorsal view
ANT=anterior BJ=body of jaw © DP=dorsal process of jaw
H = height HI = hinge of jaw L = length
MB = masticatory border MP = masticatory process
POST = posterior W = width
Radula
The radula of nudibranchs (Figures 8, 11, 14, 17, 20, 23,
26), as in most other mollusks, serves as a rasping organ
which, operated by a complicated series of muscles, moves
back and forth over the odontophore pulling food into the
pharynx. However, descriptions of the radular muscula-
ture and other functional considerations of the buccal
mass are left for future work. ;
Because the basic shape of the median and lateral teeth
is essentially the same in all the species, the reader is
referred to MacFarianp (1966) for a relatively detailed
descripion of these teeth in 4 species. The width-height
ratio for the median tooth (see Figure 6 for measurements)
referred to in this paper is not the same as that of Gar-
sTanc (1890), MacFartanp (1966) or O’DoNnocHUE
(1921); it was used because of the ease with which it
could be measured. This ratio usually decreases, some-
Vol. 12; No. 4
times markedly, from the anterior, well-worn teeth to the
posterior, unused teeth.
The denticulation and number of the median and later-
al teeth of the radula are taxonomically important. For
purposes of this paper, the lateral teeth in a transverse
row are numbered from medial to lateral, number 1 being
Figure 6
Measurements and components of a generalized Dendronotus radula
H = height LTC = lateral tooth cusp
LTD = lateral tooth denticle MTD = median tooth denticle
W = width
adjacent to the median tooth. The rows are numbered
from oldest to newest, number 1 row being the oldest.
The denticulation is a striking feature of the radula of
Dendronotus. Along the cutting edge of the triangular
cusp of the median tooth, there may be: a few relatively
large, sharp denticles; many small denticles giving a
serrulated appearance; no denticles at all; or any combi-
nation of these three. These denticles may be limited to
the proximal half of the cusp or they may extend along
its whole length, in which case the denticles generally
decrease in size towards the apex of the cusp. The denticles
of the lateral teeth are limited to the outer margin of the
sharp, curved cusp although they and the cusp are usually
absent from the innermost teeth. These denticles vary in
size, number, and shape, but there is an intraspecific con-
sistency and interspecific dissimilarity which aids in classi-
fying these nudibranchs.
Reproductive System
Dendronotus has a triaulic genital system (Figures 9, 12.
15, 18, 21, 24, 27, 28) with the penis completely separate
and the “female portion split longitudinally in an uterine
and a vaginal portion which communicate proximally and
distally’ (OpHNER, 1936, p. 1071; see also Exior, 1910;
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Page 439
GHISELIN, 1965). No attempt was made to determine
the exact relationships and functions of the various struc-
tures in this triaulic system, but the following scheme,
based on reports on related genera and families (GHISELIN,
op. cit.; ODHNER, op. cit.; THompson, 1961) is probably
reasonably accurate.
The hermaphrodite gland or ovotestis, composed of
numerous tightly packed round to pyriform lobules, pro-
duces both ova and sperm. The central rounded follicle
contains developing spermatogonia and mature sperm,
while the developing ova surround these male follicles
(MacFar.anp, 1966). A thin tube arises from this lobule
to join others, eventually forming the hermaprodite
duct through which gametes course to the ampulla
where the sperm are stored prior to copulation (GHISELIN,
1965, p. 331). There is some evidence that the sperm are
produced first and, after copulation, are stored in the
seminal receptacle of the partner until the ova ripen
(GHISELIN, op. cit., p. 343; but see also THompson, 1961,
p. 10). There is a short spermoviduct joining the am-
pulla to the bifurcation that leads to the vas deferens
and the oviduct. Possibly there is a valve at this bifur-
cation that functions much the same way as it does in
Tritonta hombergi to separate endogenous gametes and
send them along the proper ducts (THompson, op. cit.).
The sperm travel along a short duct, the proximal
portion of the vas deferens, leading to the prostate por-
tion of the vas deferens. The function of the prostate,
composed of varying numbers of large or small glandular
alveoli, or both, is poorly known (GHISELIN, op. cit., p.
331). ODHNER’s suggestion (1936, p. 1070) that the extern-
al appearance of the prostate may be species specific has
been verified in this study. The distal portion of the vas
deferens, which probably secretes some of the prostatic
secretions, continues from the prostate to the penis. This
distal portion may be short and thick (Dendronotus dalli,
Figure 15) or it may be long and tortuous (D. rufus, Fig-
ure 18). It continues to the penis tip as the convoluted, ta-
pered ejaculatory duct. The unarmed penis, enclosed
within a preputium opening to the outside, varies con-
siderably in size and shape.
The female system begins at the bifurcation of the
spermoviduct as the oviduct. This leads to the fertiliza-
tion chamber and_ subsequent parts; “albumin
gland,’ membrane gland, and mucous gland
(GHISELIN, op. cit., p. 334). Just before the oviduct enters
the fertilization chamber, an insemination duct,
which subsequently dilates to form the seminal receptacle,
arises. From the seminal receptacle, the vagina courses to
the common female cavity or vestibule into which the
“female gland mass” also opens. Near the opening of the
vagina, there is a small bursa copulatrix which may
Page 440
not be functional (GHISELIN, op. cit., p. 334; ODHNER,
op. cit., p. 1107).
Copulation in Dendronotus seems to be reciprocal, the
penis of each partner being inserted into the vagina of
the other. At present, the exact place where the sperm is
deposited is an enigma, but the length of that portion of
the penis that is inserted into the partner suggests that it
is in the seminal receptacle. The exogenous sperm is prob-
ably stored in the seminal receptacle (as indicated above)
where it may be rendered physiologically active by secre-
tions from the epithelium (THompson, op. cit.). From
here, it travels to the fertilization chamber, via the insem-
ination duct, to fertilize the endogenous ova. The zygote
follows a predetermined path through the “female gland
mass” where the primary egg capsules, nidamental layers,
and other necessary mucus coats are added prior to ovipo-
sition (GuiseLIn, 1965; THompson, 1961). The egg
masses are Type B (Hurst, 1967).
Geographical Distribution
At present, the distribution records for the species of
Dendronotus suggest that the genus is restricted to the
north temperate and arctic seas. With the exception of
one record of D. albus from Islas Los Coronados, Mexico
(Lance, pers. comm.), none have been described from the
sub-tropical or tropical seas. So far, the only record from
the seas of the southern hemisphere is 2 small specimens of
D. gracilis (Miller, pers. comm.). This type of distribution
is possibly directly related to the collection effort, but it
may represent an actual biological phenomenon caused
by so far unknown causes. Further investigations would be
desirable to clarify this situation.
As far as possible, all geographical areas where a species
of Dendronotus has been found are recorded. The records
for D. frondosus are extensive and have been condensed.
Ecology
No species description should be considered complete un-
less some indication of the species’ life history and ecology
is given. Yet, very little has been published concerning the
ecology of Dendronotus. The food preference of D. fron-
dosus has been briefly mentioned by a few authors (MIL-
LER, 1961; SWENNEN, 1961; THompson, 1964; Waters,
1966). Hurst (1967), MacFartanp (1966), MILLER
(1962), Swennen (1961), and others report briefly on
various aspects of the reproductive biology of D. frondos-
us. There are numerous rather extensive reports on the
geographical distribution of Dendronotus (Brercu, 1894;
Marcus, 1961; Opuner, 1926) giving collection sites
and occasionally depth. However, no description is given
THE VELIGER
Vol. 12; No. 4
of the hydrography or of the substratum, and both are
important in understanding the ecology of the nudibranch.
More complete ecological studies will be the subject of
another paper, but the data presented here indicate
trends and are useful both for taxonomic purposes and as
a guide for further ecological studies. The physical data
collected include depth, current action, and composition
of the bottom (rock, mud, shell-gravel, etc.). The imme-
diate substratum (hydroid, rock, etc.) of the animal at
the time of collection plus the presence (or absence) and
abundance of potential food items is noted.
Some aspects of the reproductive biology such as copu-
lation in the field and laboratory, and degree of gonadal
development are recorded. In some cases, a description of
the egg string, its location in the field, and length of time
from oviposition to hatching of the veliger is presented.
Brief descriptions are made of the feeding processes
where seen or where reported. A more detailed report of
these processes will be the subject of a future publication.
Comments concerning the possibility of cryptic or warn-
ing coloration as well as some methods of defense against,
or escape from, predators are made where appropriate.
Dendronotacea
This name was first used by OpHNER (1934) to include
a number of families and genera (OpHNER, 1936). This
suborder is characterized partly by the structure of the
liver but, more importantly, by the fact that the clavus
of the rhinophore is retractable into the rhinophore sheath
(Extot, 1910; Opuner, 1934, 1936) and by the branched
cerata on the notum. The latter are found in all the species
of the Dendronotacea to a greater or lesser degree.
DENDRONOTIDAE
Dendronotus ALDER & HANcock, 1845
At present, the family has only one genus, Dendronotus.
On the basis of a reduced branching of the cerata, crown
papillae, and veil papillae compared to Dendronotus (EI-
ot, 1910), Bercu (1863) established the genus Campaspe
to include the species C. pusilla and later, C. major BeRGH,
1886. Evior (1910) and OpHNeER (1936) conclude that
BrErGH was actually describing juvenile specimens of D.
frondosus. ODHNER (op. cit., p. 1105) states that:
“Since the genus Campaspe BERcH 1863, to which BErcH
referred two species established by himself; cannot be sepa-
rated from Dendronotus and is based on juvenile specimens
of D. frondosus ..., the family includes [a] single member the
genus Dendronotus.”
Vol. 12; No. 4
Those features characterizing the genus also necessarily
apply to the monotypic family. The following compendium
of characters has been compiled from the family or genus
diagnoses of Exior (1910), OpHNER (1936), PRruvor-
Fou (1954), Turere (1929-35).
Body limaciform; 4 - 8 pairs of arborescent cerata, one
of each pair on either side of the dorsum; cerata lacking
cnidosacs and any special gill (present in the Dotonidae) ;
relatively distinct veil supporting 2-5 pairs of more or
less branched veil papillae; clavus of rhinophore perfoli-
ate with 8 - 30 leaves; top of rhinophore sheath extended
as 4-6 more or less branched crown papillae; a lateral
papilla originating from the lateral border of the rhino-
phore stalk; anal papilla between first and second cerata
on the right side; genital openings anterior and ventral to
base of the first ceras on the right side; liver in 3 parts,
one large posterior portion and 2 smaller anterior portions;
hepatic diverticula may extend into some or all of the
cerata and rhinophores; no stomachal plates; salivary
gland long; relatively strong jaws; radula narrow to
moderately wide (6-1-6 to 21-1-21); median teeth with
smooth or denticulated cusp; lateral teeth narrow, point-
ed, usually denticulated; triaulic genital system; prostate
present; bursa copulatrix small; penis unarmed.
Dendronotus frondosus (ASCANIUS, 1774)
(Plate 63, Figure 29; Text figures 4, 7, 8, 9)
Amphitrite frondosa Ascantus, 1774, K. Norske Vidensk. Sels-
skabs Skrifter, Deel 5: 155; plt. 5, fig. 2
Doris arborescens MULLER, 1776, Zoologiae Danicae Prodro-
mus, p. 229
Doris cervina GMeEuin, 1791, Syst. Nat. per Regna Tria Nat.,
13" ed., 1 (6): 3105, no. 12
Tritonia arborescens Cuvier, 1817, Ann. Mus. 6: 434; plt. 6,
figs. 8- 10
Tritonia reynoldsii CourHouy, 1838, Boston Journ. Nat. Hist.
2 (1): 74; plt. 2, figs. 1-4
Tritonia lactea THompson, 1840, Ann. Nat. Hist. 5: 88; plt. 2,
fig. 3
Tritonia pulchella ALDER & Hancock, 1842, Ann. Mag. Nat.
Hist. 9: 33
Dendronotus arborescens (MULLER, 1776). ALDER « Hancock,
1845, Ray Soc. Monogr., Fam. 3, plt. 3, prt. 1
Am phitritidea fabricti “Beck 1847.” Morcu, Grénland
(Rink) (Prodr. Fauna Moll. Gronl., p. 6), 1857; in
synonymy [this reference has been taken directly from
TREDALE & O’DonocHuE, 1923]
Dendronotus luteolus LaFont, 1871, Act. Soc. Linn. Bor-
deaux, 28: 287; plt. 17, fig. 1
Dendronotus purpureus BercH, 1879, Proc. Acad. Nat. Sci.
Philadelphia, art. 5: 145 - 150; pit. 1, figs. 18 - 20; plt.
3, figs. 7 - 12
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Page 441
Dendronotus purpureus var. aurantiaca FriELE, 1879, Jahrb.
Deutsch. Malakozool. Gesellsch. 6: (page unknown)
[reference unavailable]
Campaspe pusilla Bercu, 1863, Naturh. Tidsskr., 3 R., 1:
471-478; plt. 12, figs. 28 - 35
Campaspe major Bercu, 1886, Bidjr. Dierk. 13: 21 - 24; plt. 1,
figs. 23 - 26; plt. 2, figs. 1 - 11
Dendronotus frondosus (Ascantus, 1774). Plymouth Marine
Invert. Fauna 1904. Journ. Marine Biol. Assoc. U. K.
7: 282
Dendronotus lacteus (THompson, 1840). Extor, 1910, Ray
Soc. Monogr. Suppl., Fam. 2, genus 1, p. 161
Dendronotus venustus MacFarLanp, 1966, Mem. Calif. Acad.
Sci. 6: 271 - 275; plt. 40, fig. 2; plt. 46, figs. 9 - 12; plt.
47, figs. 1-2; plt. 49, fig. 6; plt. 50, fig. 3; plt. 52,
figs. 3-6
Taxonomic Remarks
The above references are only part of a fairly extensive,
and often conflicting, literature concerned mainly with
taxonomy, morphology, and geographical distribution of
Dendronotus frondosus.
The description by Ascanius (1774) is sufficient to
establish that Amphitrite frondosa is the same animal as
Doris arborescens Mi.iER, 1776. Thus the name Amphi-
trite frondosa takes priority. However, the generic name is
unavailable because it is used by O. F MUxier in 1771 for
a genus of polychaetes. Because A. frondosa is not the type
species for either the genus Doris or Tritonia, nor is it a
member of either of these genera, the species must be
placed in the next available genus, Dendronotus, estab-
lished by ALDER & Hancock (1845). They distinguished
Dendronotus from Tritonia on the basis of rhinophore
structure, the lack of a sub-pallial ridge on the dorsum,
free arborescent cerata, and the structure of the liver.
However, they used the species name arborescens appar-
ently unaware of the existence of frondosa. The first time
Dendronotus frondosus was used appears to be in the
Plymouth Marine Invertebrate Fauna (1904). Probably
Euiot was responsible for this change (see Exiot, 1910)
although no reference is made to him. Subsequent authors
have used both D. frondosus and D. arborescens, but the
correct name is Dendronotus frondosus (Ascantus, 1774).
Biologists of the 18 and 19% centuries added to the
confusion by describing a number of “new species” (Dend-
ronotus luteolus, D. purpureus, D. lacteus) as well as a
“new genus,’ Campaspe. All of these are almost surely
synonyms of D. frondosus.
OpuHNER (1936) used Dendronotus frondosus as a
catch-all for all the previously described species except
D. robustus and D. iris. He maintained that D. frondosus
was a highly polymorphic species and that the other de-
scribed species were well within the limits of variability
Page 442
to be expected from this polymorphic species (see under
D. dalli and D. rufus for further discussion) .
Dendronotus venustus represents a case of splitting.
Except for color, this species is very similar to D. fron-
dosus. I have collected a few specimens which, in color,
intergrade between these two nominal species, particularly
with reference to the white patches found between the
cerata by MacFartanp. In his field notes (deposited at
the California Academy of Sciences), MacFarLanp de-
scribes color phases of D. venustus which are identical
with D. frondosus. Lance (pers. comm.) is also of the
opinion that D. venustus is a synonym of D. frondosus.
The descriptions of Dendronotus frondosus by Marcus
(1961) and MacFartanp (1966) confuse 3 or 4, and 3,
distinct species, respectively.
Body Dimensiens, Texture, and Apertures
The limaciform, laterally compressed body is moderately
“heavy.” The rounded dorsum tapers posteriorly to a
long, sharply pointed tail and merges laterally with the
vertical sides.
The largest animal collected was 4.2 cm L while the
largest reported are 5.5-6.0cm (BERGH, 1879, 1894),
9.8cm L (SwENNEN, 1961), and 11.5cm L (MacGinimE,
1959). The smallest animal I collected was 0.2cm L.
Most, however, have been 1.0 - 3.0cm L with only a few
being longer than 3.5 cm.
The anus is borne on a distinct papilla, often capped
with a yellow pigment, and located halfway between the
first and second right cerata.
The genital apertures open externally about 3 of the way
up the right side of the body, just anterior to the base of
the first ceras.
The cardiac prominence is large and often rises 4-5
mm higher than the dorsum. The body is covered with
bluntly conical papillae of various sizes that are usually
tallest on the cardiac prominence. A distinctive charac-
teristic of this species is the yellow or white pigment that
caps these papillae; only rarely is it missing.
Foot
The long, relatively narrow foot is bluntly rounded ante-
riorly and tapers to a short, pointed tail. The sole is white
and translucent.
The edge of the foot flares along its entire length when
the animal is crawling on a smooth surface. It is able to
adhere more firmly than other species to smooth surfaces.
When on a hydroid, the edges of the foot wrap around the
stalk allowing the nudibranch to adhere firmly to the
hydroid.
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Vol. 12; No. 4
Color
The translucent, gray-white ground color of the animal is
overlaid with varying concentrations of brown, red-brown,
yellow, and white pigments in assorted patterns (Plate
63, Figure 29). To the naked eye, some of the animals
have a reddish-brown hue, streaked and marbled with
brown and spotted with yellow and white while others
are very pale tan with numerous yellow spots. A few,
particularly those from deeper water (more than 25m),
lack all pigment and appear white.
In about 50% of the animals, the body was brown with
a sparse scattering of white and yellow spots, but the distal
portions of all the body processes were devoid of brown
and appeared transparent or yellowish. A few animals
were devoid of light pigments, while a few others were
devoid of any brown but were liberally covered with yel-
low or white or both (see ALDER & HANcock for colora-
tion and pattern).
The pinkish-brown hepatic diverticula and liver are
clearly visible through the body wall as is the white
hermaphrodite gland.
Cerata
The 5-7 (4-8) pairs of tall, erect cerata are arranged
at posteriorly decreasing intervals. The height and degree
of branching also decreases posteriorly. In a few speci-
mens, there are | - 4 small, unbranched, unpaired cerata
on the posterior end.
The cerata are more arborescent than in the new species
of Dendronotus, but less so than in D. rufus or D. iris. The
main branches, quite tall and slender, give rise to numerous
relatively long secondary and tertiary branches that end
in pointed transparent tips.
The pattern of main branches is typically “fan-shaped”
(Figure 4a) with 3 main branches in the first 3 pairs, 2 - 3
in the 4" pair, 1 - 2 in the 5", and one in the rest. In those
pairs with 3 main branches, the medial is the tallest while
no. 2 branch is about # as long and arises just above the
lateral. The lateral branch, arising right at the base of the
ceras, is about 4 to 2 as long as the medial and diminishes
in size from the anterior to the posterior cerata.
The hepatic diverticula appear to be present in the
anterior 4-5 pairs of cerata, usually in the medial 1 - 2
main branches as well as the rhinophores. These divertic-
ula in the first pair of cerata and the rhinophores originate
from the anterior lobes of the liver and the rest arise from
the posterior lobe.
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Page 443
Rhinophore and Clavus
The moderately branched rhinophore is about equal in
height to the first pair of cerata.
The lateral papilla, arising between a quarter and half
way up the stalk, is relatively long, often extending as
high as or higher than the clavus. The degree of branching
is variable; some are simple papillae, most have a few
short, simple secondary branches, and a few are very
branched, much like the cerata.
The sub-conical perfoliate clavus bears 8 - 12 shallow-
cut leaves that alternate in width. There are a few reports
of up to 20 - 25 leaves in larger animals (Bercu, 1894,
1900) and I have seen as few as 5 - 6 in smaller ones. The
coloration of the clavus is similar to that of the rest of
the body.
There are usually 5 (4-6) long, stout crown papillae
having little or no secondary branching. The postero-me-
dial papilla may be 1-3 as long as the 3 anterior
papillae which are usually about equal in length. The
postero-lateral may be about twice as long as the anterior
ones.
Head, Lips, and Veil
In size, number, and branching, the veil papillae seem
to come midway between the condition found in Dendro-
notus rufus and D. dalli, and that of D. albus.
There are 4 pairs of relatively stout, branched veil
papillae. On the larger pairs, the branching may be ex-
tensive, much like the cerata, or it may be confined to a
few small papillae. The smaller pairs are often unbranched,
sometimes have a few small papillae, or, rarely, they
may be extensively branched. The medial pair (no. 1) is
the longest, no. 2 is the shortest at about } to 4 as long,
no. 3 is slightly shorter than no. 1, and no. 4 is about 3 as
long as no. 1.
There are 2-4 (0-8) long, thin, simple lip papillae
located lateral and posterior to the plicated lips. These
papillae are sensitive to touch and retract almost com-
pletely when stimulated mechanically.
Jaws
The body of the jaw (Figure 7) is a translucent dingy-
yellow which becomes dark brown, almost black on the
masticatory process, hinge, and proximal portion of the
dorsal process. The body is a shallowly convex, almost
oblong structure about 24> as long as wide with a mod-
erately convex posterior end.
The slightly curved, rather wide dorsal process is in-
clined posteriorly at 55° - 60° from the long axis of the
Figure 7
Dendronotus frondosus jaw
B. Ventral view C. Outside lateral view
D. Inside lateral view
A. Dorsal view
body. It is about 0.41 as long as the body of the jaw
and has a shallow dorsal groove. A thin, strongly convex
lateral expansion joins the proximal 3 of the process to
the body. The moderately long, curved masticatory pro-
cess, often sharply hooked at the free end, is joined by an
almost flat, strong expansion to the body. A small number
of relatively large, black denticles adorns the masticatory
margin.
Radula
The radula (Figure 8), described a number of times
(AtpEeR &« Hancock, 1845; Bercu, 1879, 1894, 1900;
Meyer « MoOsius, 1865; O’DonocHuE, 1921), varies
widely in the number of rows of teeth present, but this
appears to be directly correlated with the length of the
animal. The radula formula, from the literature, is
29 -49(7 - 14:1-7-14) and from those I examined (10
specimens), it is 33-48(7-11-1-7-11). Bercu (1894)
and VoLopcHENKOo (1955) report that they found golden-
yellow teeth in their specimens, but all those I examined
were colorless.
The cusp of the median tooth, about 1.4 - 1.7 as wide
as high, comes to a relatively sharp point. There are 7 to
15 large, strong, sharp denticles on the sides of the cusp,
usually becoming smaller towards the apex. In the ante-
rior teeth, there are fewer denticles and they tend to
Page 444
disappear near the apex while the posterior teeth have
more denticles extending along the whole side. The lateral
edge of the denticles continues down the dorsal side of
the cusp as a deep furrow.
The elongate, relatively wide lateral teeth bear long,
curved cusps inclined 10° to 20° toward the midline.
They are usually rather longer than shown in Figure 8.
Figure 8
Dendronotus frondosus radula
Rows 25 to 27 in a radula with 45 rows of teeth
On the innermost 1 - 3 teeth, the cusp is often no longer
than the denticles while in the outermost 1 - 2, the cusp
may be missing or rudimentary. The size is about equal
in the rest. From anterior to posterior, within a single
radula, the cusps tend to become longer and fewer are
broken off.
All the teeth, except the outer 1-2, bear 2-5 (0-7)
long, strong, sharp, regularly spaced denticles on the
lateral margin of the cusp.
Reproductive System
The large, white hermaphrodite gland gives rise to a
long hermaphrodite duct that widens suddenly into a wide
ampulla curled up like a doughnut (Figure 9). The long,
narrow spermoviduct arises from the other end of the
ampulla but runs alongside the hermaphrodite duct for
a short distance. The proximal portion of the vas defer-
ens and the oviduct are both short, translucent and quite
narrow.
OpHNER (1936) reports -and figures the prostate as
being “. . . a circular disc composed of numerous scattered
vesiculae [= alveoli] (fig. 39a), in many circles ...”
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Vol. 12; No. 4
Figure 9
Reproductive system of Dendronotus frondosus (exploded view)
AMP = ampulla BC = bursa copulatrix
DVD = distal vas deferens FC = fertilization chamber
FGA = female genital aperture FGM = (to) female gland mass
HD = hermaphrodite duct ID = insemination duct
OV = oviduct P= preputium PE = penis PR = prostate
PVD = proximal vas deferens SO = spermoviduct
SR = seminal receptacle VA = vagina VE = vestibule
However, I rarely saw more than 12 and usually only 5 - 8
large, ovoid alveoli arranged as a disc concentric with
the vas deferens. From the prostate arises a very narrow,
long, convoluted, almost untapered distal portion of the
vas deferens. It enters the penis to become a much con-
voluted ejaculatory duct. The very long, thin penis, tapered
to a point and coiled up in the preputium, has a pleated
base.
The long, convoluted vagina empties into the large,
sessile, pear-shaped seminal receptacle. The distal end of
the vagina opens into the vestibule quite near the external
orifice. A small, stalked bursa copulatrix is located about
3 of the length of the vagina away from the vestibule.
The insemination duct is moderately long and convoluted.
The reproductive system described and figured by Mac-
FarLanp (1966) for Dendronotus venustus agrees closely
with that of D. frondosus, further evidence that D. ven-
ustus should be considered a synonym of D. frondosus.
Geographical Distribution
Dendronotus frondosus, a north circumpolar species, has
been reported from Arctic seas and the northern Atlantic
and Pacific Oceans. More exact locations include the
coast of Norway (OpHNER, 1926, 1939), Point Barrow,
Vol. 12; No. 4
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Page 445
the Bering Sea, the east coast of North America from
Labrador to Cape Cod, Greenland, the Faroes, the Shet-
lands, Spitzbergen, and the western Arctic (MacGint-
TIE, 1959), the eastern Atlantic south to the Bay of
Biscay (Bercu, 1894), and the Baltic Sea as far as
Kiel Bay (SwENNEN, 1961). It has also been found on
the northern Asiatic coast (Basa, 1957; VoLopcHENKO,
1955) and along the Eastern Pacific Coast from Alaska
to Southern California (Lance, pers. comm.; unpublished
observation). Kerry Clark (pers. comm.) has collected a
number of specimens from the coast of Connecticut.
Ecology
Despite the number of descriptions of the species, there
has been surprisingly little published about its biology.
Brief mention has been made of its food and substratum
preferences, and the reproductive cycle (Murtter, 1961,
1962; SweNNEN, 1961; THompson, 1964; WarTERs,
1966).
Locally, the species has been collected from April to the
beginning of December from floating wharves in Friday
Harbor and from June to March in the sub-tidal down
to 40m. SwENNEN (1961) reports finding Dendronotus
frondosus all year long at Den Helder. Other reports
(Miter, 1962) would indicate D. frondosus are probab-
ly present as adults, in greater or lesser numbers, through-
out the year. Mr. K. Clark (pers. comm.) has found spe-
cimens in May, but not in summer, in Connecticut.
Dendronotus frondosus ranges from the inter-tidal to
400m-+ (SwENNEN, 1961) and is present on rocky as
well as muddy bottoms and styrofoam floats, where
numerous hydroid colonies are present.
Dendronotus frondosus appears to be a feeding spe-
cialist on hydroids, but is flexible about what species it
will eat. It has been reported to feed on Tubularia indivi-
sa and Sertularia cupressina (SWENNEN, 1961), Syn-
coryne eximia, Bougainuillia glorietta, and Obelia com-
missuralis (WaTERS, 1966), Tuwbularia larynx (BRAAMS
& GEELEN, 1953), Dynamena pumila, Hydrallmania
falcata and Sertularia argentea (Mituer, 1961),and Tubu-
laria crocea (K. Clark, pers. comm.). There is also some
evidence that the younger animals prefer one species of
hydroid while the adults prefer another (SwENNEN, op.
cit.). Noting this, MiLter (1961, p. 105) says:
“... small specimens (1- 18mm) of Dendronotus frondosus
feed on both gymnoblastic and calyptoblastic hydroids, large
specimens (14-100mm) only on gymnoblasts (Tubularia
spp.). The large polyps of Tubularia probably provide a fairly
large volume of food in a form which can be grazed easily
and rapidly, supplying a quantity sufficient for the needs of
such a large nudibranch.”
However, preliminary laboratory and field observations
(Waters, 1966; author’s unpublished data) suggest that
this is not so locally and that adult D. frondosus prefer
thecate hydroids like Obelia to athecates like Tubularia.
The mechanism of feeding has been briefly described
(Waters, 1966), and some additional information has
since been obtained, particularly pertaining to predation
on Obelia commissuralis, When the veil and lip papillae
come in contact with a hydranth, the nudibranch imme-
diately retracts these structures, and then begins to search
about for the prey. This withdrawal, presumably in res-
ponse to the nematocysts of the hydranth, may occur 2 to
3 times, but eventually the animal envelops the whole
hydranth and, using the jaws and radula, chops it off.
Occasionally the nudibranch will attack just proximal to
the hydranth. In this case, it uses the same method as
when the coenosarc is eaten (WATERS, op. cit.). The
nudibranch appears to use the outer lips and lip papillae
to align the body and opening of the jaws parallel to the
stalk. Achieving this, it grips the perisarc firmly with the
outer lips, the jaws are protruded, and working trans-
versely to the stem’s long axis, they cut a hole in the
perisarc. The radula is apparently extended through this
hole and proceeds to rasp out the coenosarc (or the
hydranth). In a relatively short time, one nudibranch is
able to clean out a large hydroid colony.
There are conflicting reports about the reproductive
activity of the species. SWENNEN (1961)reports it as
being sexually mature all year except February and No-
vember, and spawning from April to August and again
in January. Locally, spawn and mature animals were
seen from April to October. Clark (pers. comm.) found
egg masses during early May in Connecticut. THomPsoNn
(1964) and Mitter (1962) indicate that Dendronotus
frondosus may live for 2 years, breeding in both years.
They also suggest that there may be more than one
generation a year, but SWENNEN (op. cit.) feels that
most only live one year. It is possible that a few animals
do live for 2 years, not having spawned the first year,
but most probably mate, spawn, and die within a year.
The nidosome is described by Hurst (1967, p. 264)
as “... an untidy coil varying from pale to dark pink.”
I have also seen a number of white ones in relatively
neat coils. There was usually only one egg per capsule,
and the eggs took from 13 to 16 days to hatch.
It is hard to say how cryptic the coloration of this
nudibranch is because the background may be so variable.
On hydroids, it would likely not show even in its lighter
brown color forms although the white, deepwater ani-
mals are quite visible to a diver. There are no other
obvious means of defense although it may have some
Page 446
epidermal secretions (THomMpson, 1960b; Epmunps,
1966).
Dendronotus frondosus is an able swimmer, but the
actual distance covered is usually small unless a current
is present. Mechanical stimuli such as poking or pinching
with forceps are effective in eliciting the swimming be-
havior. So far, no potential predator has been demon-
strated to elicit swimming although VoLopCHENKO
(1955) reports that this species is preyed upon by fish.
Dendronotus iris Cooper, 1863
(Plate 63, Figure 30; Text figures 4 - 6, 10 - 12)
Dendronotus iris Cooper, 1863, Proc. Calif. Acad. Nat. Sci.
3: 59
Dendronotus giganteus O’DoNocHuE, 1921, Trans. Roy. Ca-
nad. Inst. 13 (1): 187 - 190; plt. 4, fig. 47; plt. 5, figs.
57 - 59
Taxonomic Remarks
Coorper’s brief description, based on color, is adequate to
establish Dendronotus iris as a valid species name. He
states (p. 59) that “this species seems more variable in
color than other nudibranchiata of this coast, but I [see]
no reason for considering [it] more than one species” and
indicates that a slight color variation of it may be found
in Puget Sound.
O’DonocHuE (1921) recognizes that color is quite vari-
able in the nudibranchs, even between Nanaimo, B. C.
and California, but he neglects this when he describes
Dendronotus giganteus. He makes no reference to Coo-
PER’S paper.
OpHNER (1936, pp. 1107 - 1108) recognized O’Dono-
GHUE'’ error and declared Dendronotus giganteus a syno-
nym of D. iris. Subsequent authors, with the exception of
SmitH « Gorpon (1948), have retained D. iris as the
proper name.
Body Dimensions, Texture, and Apertures
The limaciform body, the “heaviest” of all the local
species of Dendonotus, tapers abruptly to a bluntly
pointed tail. A well-rounded dorsum merges into vertical
sides, the only demarcation being the 2 rows of cerata.
The largest animal found was 29.0cm L & 6.4cm H
< 8.9cm W and displaced 1100 cc of water. O’Dono-
GHUE (1921) claims a preserved specimen, measuring
21.0cm L & 8.4cm H X5.5cm W, may have been 26cm
L < 10.0cm H & 6.5cm W when alive. Most of the
animals (many preserved) were 6.5 - 12.0 (3.0- 29.0) cm
L X 1.5-2.0 (0.5-5.0) cm H X 1.0-2.2 (0.5-4.0) cm
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Vol. 12; No. 4
W.A few representative animals were: 8.0cm L & 1.5cm
H X 1.5cm W (live); 65cm L X .1.5cm H X'1.0cm
W (live); 9.0cem L & 3.5cmH X 2.5 cm W (preserved);
3.0cm L X0.5cm HX 0.5cm W (preserved).
The anus is on a distinct, truncate papilla located half
way along the imaginary line joining the bases of the
first and second right cerata. Often, an opaque white
line marks the edge of the papilla and the plicated lips,
surrounding the anal opening, are usually white.
The distinct genital openings are located about 2 the
way up the right side, below or immediately anterior to
the base of the first right ceras. Sometimes, an opaque
white line marks the edge of the genital openings. Very
often, in preserved specimens, the penis is extruded.
The cardiac prominence is usually not visible except
as a very slight bulge. The body was smooth in all the
specimens examined.
Foot
The long, quite wide foot is bluntly rounded anteriorly
and terminates in a short, bluntly-rounded tail. When
crawling, the foot often flares out considerably beyond the
plane of the sides of the body, contrary to what MacFar-
LAND (1966, p. 258) says. In one 8cm L animal, the
foot was flared out 1.5cm on either side, giving the foot
a total width of 5.5 cm.
A useful, but not infallible, diagnostic character of the
species is the narrow, opaque, dead-white line that edges
the dorsal margin of the foot. The sole is white in the
“gray” forms and a light orange in the “orange-red”
forms.
The expansiveness of the foot may be an adaptation to
crawling across the soft muddy bottom that this species
inhabits, the greater surface area allowing for a better
grip on the unstable substratum. It is almost non-func-
tional as a grasping organ, and apparently is non-func-
tional in helping the animal to maintain its position in
a fast current (which it probably encounters only rarely
in nature).
Color
Most specimens (23 of 30) were of the “gray” form,
while 7 were of the “orange-red” form. In the “gray”
form, the ground color varies from a clean, translucent,
gray-white through yellow-brown to a dark muddy brown
(Plate 63, Figure 30). The ground color is lightest on
the sides, becoming darker on the dorsum and bases of
the dorsal processes (cerata and rhinophores) and very
dark towards the distal ends of the cerata, crown papillae,
lateral papillae, and veil papillae. At the ends of the
Vol. 12; No. 4
THE VELIGER
Page 447
smaller branches of these processes is a variety of colors:
metallic orange (1 animal), dark purple (11 animals), a
mixture of chrome-yellow and purple (9 animals) with
small amounts of white found in most animals.
In the “orange-red” forms, the ground color is a beauti-
ful “orange-red.” Again, this becomes darker on the cera-
ta, rhinophores and veil papillae, but finally gives way,
at the ends of the appendages, to purple (2 animals),
metallic orange (3 animals), or a mixture of the two
(2 animals).
Only one animal had the white tips on the cerata with
a sub-terminal orange ring described by Cooper (1863).
All the intergradations between the two main color forms
exist.
The lip papillae are normally tipped with chrome yel-
low or are unpigmented, but 2 specimens had metallic
orange in place of the yellow.
Cerata
There are 4-6 (3-8) pairs of stout, but very tall and
dendriform cerata, the last pair being about half as tall
as the first. Arranged at decreasing intervals on the dorso-
lateral margins of the body, they begin about 4 of the way
back from the anterior end. Nearly all the specimens
have 2 (1-4) small, branched, unpaired papillae on the
posterior end.
The cerata are not firm and erect as they are in the
less dendritic forms, but are instead quite supple and
extensible. Because of these traits, and the fact that they
are usually badly mutilated in the dredge hauls, it is
often difficult to determine the number of main branches
in a ceras.
The very thick, basal portion of the ceras divides al-
most immediately to give rise to a number of tall, thick
main branches arranged in a “fan-shaped” pattern (Fig-
ure 4a). There are 3 (2-5) main branches in the first
pair (no. 1), 3 (2-4) inno, 2, 2-3 inno. 3, 2 (1-3) in
no. 4, 1 (1-3) inno. 5, 1 (0-2) inno. 6, and 1 in nos. 7
and 8. These subdivide into secondary branches that may
be nearly as long as the main ones while the tertiary
branches are only slightly shorter. The secondary and
tertiary branches end in small, delicate tufts, giving the
animal a very bushy appearance when viewed from
above. The medial branch is the longest (up to 5-7cm
ina 15cm L animal) while the lateral is the shortest.
According to MacFarLanp (1966) only the first pair
of cerata and the rhinophores have “ceratal cores” or
hepatic diverticula, arising from the anterior lobes of the
liver. OpHNER (1936) states that the diverticula are
ramified throughout the cerata as shown by ALDER &
Hancock (1845), but neglects to mention in how many
pairs this occurs. In 10 animals examined, I always found
the situation described by MacFarLanp.
Rhinophore and Clavus
The rhinophore resembles the cerata with its very thick
stalk and the extensive branching of the lateral and crown
papillae. At the base, the lateral papilla is thick but it
soon divides into a number of smaller secondary branches
which in turn branch and terminate in small, delicate
tufts. Arranged vertically on the posterior border of the
rhinophore stalk are 4 (2-6) small, branched papillae
found only in this species.
The inconspicuous, muddy brown, conical clavus is per-
foliated with about 25 (15-31) leaves alternating in
breadth. The top 2 - 4 and bottom 4 - 6 leaves are usually
small and equal. Those inbetween alternate more or less
regularly with the wider leaves almost completely cover-
ing the narrower ones.
The 5 (2-5) crown papillae are relatively longer and
more branched than in most species except Dendronotus
rufus. The branching is not as neat as in D. rufus but
tends to occur at any point from the base to the tip,
making it hard to ascertain the actual number of crown
papillae in many cases. The postero-medial is the longest,
being 2-4 longer than the 3 almost equal, anterior
papillae. The postero-lateral may be from 1} - 3 longer
than the anterior papillae.
Head, Lips, and Veil
On the indistinct veil, there are almost always 3 (2 - 4)
pairs of stout veil papillae varying considerably in size
and degree of branching. However, they are generally
sparsely branched and short, relative to the animal’s size.
Contrary to what MacFartanp (1966) shows in his
drawing compiled from a number of specimens, I found
that the medial pair was usually smallest and the lateral
pair (no. 3) the largest (from 14 0 2 longer than no.
1). Pair no. 3 is divided almost from the base into a
smaller, sparsely branched lateral ramus and a much
longer (2), more branched medial ramus. Often, the
veil papillae were all very small and simple, making
them indistinguishable from the lip papillae except by
position.
There are many lip papillae (20-40, depending on
the size of the specimen) arranged in 3 - 4 irregularly cres-
centic rows around the mouth and lips. The most anterior
row, immediately ventral to the veil papillae, consists
of 6 (5-8) simple to slightly branched papillae about
equal in length to the shortest veil papillae. In the second
row, there are 7 (6-12) simple papillae about 4 to 2
Page 448
the size of the preceding ones. The most ventral “row”
of 10-25 small, simple papillae and tubercles is very
irregularly arranged around the mouth.
Jaws
The jaws of Dendronotus iris (Figure 10) are the largest
and heaviest seen in any of the local species of Dendro-
notus. In the whole genus, only D. robustus may have
heavier, albeit relatively shorter, jaws. MacFarLANnp
(1966) found in a 9.1cm L preserved specimen that the
jaws were 1.72cm L X 0.75cm W (see Figure 5 for
measurements ). In 5 specimens that I examined, the sizes
were:
Dendronotus iris
length of animal _lengthof jaw height of jaw _—- width of jaw
(measurements in centimeters)
13.0 2.8 0.95 0.85
7.0 1.5 0.65
8.0 1.6 0.65
10.0 1.8 0.70
4.0 1.1 0.35
The total length of the animal body is only about 5X
that of the jaw.
Figure 10
Dendronotus iris jaw
B. Ventral view C. Inside lateral view
D. Outside lateral view
A. Dorsal view
The jaws are a deep reddish brown on the masticatory
process and hinge, fading to a light yellow along the
dorsal process and body. The masticatory denticles may
be almost black.
The body of the jaw is an elongate, mytiloid shape,
widest at the posterior end. The length of the body is
slightly less than 3 the maximum width which occurs
in the posterior third. The strong dorsal process, weakly
curved, is inclined posteriorly at about 40° to the longi-
THE VELIGER
Vol. 12; No. 4
tudinal axis of the body, and is joined to the body by a
strong, convex expansion for over half of the length of
the process. The dorsal process is about a third (or slightly
more) as long as the body. The strong, arched mastica-
tory process is joined to the body by another thick, but
weakly concave expansion. There are about 85 - 100 large
denticles on the masticatory border.
A feature not seen in any other Dendronotus is the very
long, convoluted esophagus. Its function is not known at
present, although it may be related to the fact that the
animal can extrude the whole buccal mass through the
lips when feeding (moribund animals usually extrude
the “proboscis’’) .
For a more complete description of the jaw, the
reader is referred to MacFarLanp (1966, pp. 259 - 260).
Radula
The radula (Figure 11) is most unlike that of any other
Dendronotus. The radula formula, according to MacFar-
LAND (1966) and O’DonocHUE (1921) is:
34-46 (11-20-1-11-20) and 35-40 (12 - 16- 1-12-16),
respectively. In 9 specimens, I found it to be:
41-61 (11-21-1-11-21), giving a maximum range of
seal (uilozil eile hil Bil).
The tall sharp-pointed cusp of the medial tooth is only
slightly higher than wide (Figure 6) and bears on the
concave margins 11 (9-18) strongly developed, pointed
denticles. These denticles, more prominent than in any
other species, are slightly concave on the medial edge. The
lateral margin of the denticle continues anteriorly on the
tooth as a deep furrow. Toward the apex of the cusp,
particularly on teeth from the posterior half of the radula,
the denticles decrease in size, number, and regularity. For
example, in one radula with 58 teeth, there were 7
large, regularly spaced denticles on the first medial tooth,
Figure 11
Dendronotus iris radula
Rows 22 to 24 in a radula with 43 rows of teeth
Vol. 12; No. 4
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Page 449
9 on the 25", 12 on the 35" and only 8 irregular ones on
the 524.
The lateral teeth become more numerous from the
oldest rows to the youngest. The oldest 1 - 3 rows have no
lateral teeth, rows 2-5 have 3-10 teeth, and by about
row 8, the main battery of teeth is present with only 1 - 3
more to be added toward the posterior end. Usually,
there are no denticles on the laterals but occasionally,
the no. 1 tooth in the first 10 - 15 rows will be serrulated
or even bear 1 - 8 short, sharp denticles. Rarely, the no. 2
lateral in the first few rows will also be serrulated or
bear a few tiny denticles.
The typical lateral tooth consists of a flat, elongated
base upon which there is a gently tapering, blunt-pointed,
stout cusp inclined at about 20° toward the midline of the
radula. The outermost 3 - 4 lateral teeth decrease in size,
the most lateral one being a narrow, flat plate with a
short spine. The innermost lateral has a rod-like base
that terminates in a short, stout cusp.
Reproductive System
The thin hermaphrodite duct joins the large, white, lob-
ulated hermaphrodite gland to the very long, tortuous
ampulla which is relatively the longest found in the genus
(Figure 12). The ampulla merges indistinctly with the
spermoviduct which bifurcates, giving rise to a short prox-
imal portion of the vas deferens and relatively long,
thin-walled, almost transparent oviduct.
The prostate is made up of alveoli surrounding and con-
cealing more than # of the total length of the looped vas
deferens. The short, narrow, muscular distal portion of
the vas deferens enters the base of the penis where it con-
tinues as the coiled ejaculatory duct. The long penis,
sheathed in a relatively thick-walled preputium, is very
thick throughout its whole length; it is the largest in
the genus.
The short, wide, muscular vagina terminates in an in-
distinct, somewhat irregularly shaped seminal receptacle.
Located on the distal third of the vagina is the bursa cop-
ulatrix atop a very long, thin stalk. A long, convoluted
insemination duct, much narrower than the vagina, joins
the seminal receptacle to the fertilization chamber via
the oviduct.
OpuHNER (1936, figure 40), in a poor diagram of the
Dendronotus iris genital system, has misplaced the in-
semination duct, a continuation of what he calls the vesi-
cula seminalis (v.s.), and the spermoviduct, a continua-
tion of what he calls the ampulla (f.). The spermoviduct
should terminate between the prostate and the insemina-
tion duct.
AMP 1.0 cm
HD |
Figure 12
Reproductive system of Dendronotus iris (exploded view)
AMP = ampulla BG = bursa copulatrix
DVD = distal vas deferens ED = ejaculatory duct
FC = fertilization chamber FGA = female genital aperture
FGM = (to) female gland mass HD = hermaphrodite duct
ID = insemination duct OV = oviduct P = preputium
PE = penis PR = prostate PVD = proximal vas deferens
SO = spermoviduct SR = seminal receptacle VA = vagina
VE = vestibule
Geographical Distribution
Dendronotus iris has usually been collected subtidally
from 7 m at Nanaimo, B. C. to about 200 m off Unalaska
in the Aleutian Islands. The southern limit at present is
the Coronados Islands in Mexico (Lance, pers. comm.).
It has been recorded from Nanaimo, B. C. (O’DonocHUE,
1921) ; the Queen Charlotte Islands, B. C. (D. B. Quayle,
pers. comm.) ; Puget Sound (Cooprr, 1863) ; and numer-
ous areas in Califomia (SmirH « Gorpon, 1948; G.E.
MacGinitie, pers. comm.). AGERSBorG (1922) reports
one from the shore of Shaw Island and another from logs
near the Puget Sound Biological Station, both in the San
Juan Archipelago.
Page 450
Ecology
This species has been obtained by dredging in a number
of places around the San Juan Islands (see Materials and
Methods, above) where it is apparently present year
round. I collected 15 more while SCUBA diving at Na-
naimo.
The animals are almost always collected from muddy
bottoms where there is little current action. This type of
bottom often supports a dense population of Cerianthus
sp., the burrowing anthozoan that is the principal food
of Dendronotus iris; in fact, the available evidence would
suggest that it is the sole food of this nudibranch.
Dendronotus iris displays a specialized feeding behavior.
How it locates the Cerianthus is unknown, but, upon con-
tact with the anemone’s tube (which extends as much as
15 cm above the substratum), D. iris begins to crawl up
the tube. The veil papillae are extended and the head
moves slowly back and forth through about a 120° arc.
When D. iris contacts a Cerianthus tentacle, the nudi-
branch immediately withdraws. Again the veil papillae
are extended and the head moves back and forth in ever
decreasing arcs until D. iris has “homed in” on the ten-
tacle. Dendronotus iris then lifts the anterior third of the
body free of the tube, lunges forward, at the same time
extruding the buccal mass through and past the mouth.
The Cerianthus tentacles are grasped with the jaws and
chopped off whole by the radula. This attack must be
and is done rapidly as the Cerianthus is able to withdraw
quickly. Withdrawal seems to be a poor defense; smaller
D. tris apparently can climb down the tube or are pulled
down the tube when the attacked Cerianthus retracts.
I have found whole, undigested Cerianthus tentacles
in the stomachs of many specimens as well as multitudes
of nematocysts in the intestine and fecal pellets; no other
prey species were found. MacFartanp (1922) reports
finding fragments of a nemertean in one animal. AcERS-
BORG (1922) tried to feed many things to a Dendronotus
giganteus (= D. iris), but it was only interested in the
tentacles from an unnamed coelenterate which were
mouthed, but not eaten.
Very little information is available concerning reproduc-
tion in Dendronotus iris. In the laboratory, between Feb-
ruary 7 and 13, 1967, 3 white egg masses were deposited
in a large rounded bundle. There were 45 - 50 and 31 - 40
eggs per capsule. The veligers hatched after 13-15, 16,
and 17 days from the 3 egg masses. In June, 1967, 3 more
egg masses with up to 100 eggs per capsule were laid by 3
very large animals (28.7, 25.4, and 20.3cm L). The same
animals were copulating and laying eggs when collected
at Clarke Rock, Nanaimo. In June, 1968, 8 egg masses
were seen on the tubes of Cerianthus sp. at the Monterey
Breakwater, Monterey, California.
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Vol. 12; No. 4
There is some suggestion that this species, particularly
the gray form, is cryptic in its natural habitat although
first-hand observation indicates that it can be spotted
fairly easily by a diver. Predators, probably fish, must
occasionally detect Dendronotus iris, either by sight or
smell, and attempt to eat the nudibranch. Evidence for
this is found in the numerous D. iris collected with parts
of the dorsal appendages missing.
The cerata (or portions thereof) are rather easily
broken off with rough handling. Upon severance, these
parts move quite naturally and even “swim” to some
degree. This may be a protective measure, serving as a
decoy while the nudibranch makes its escape.
AcErspore (1922) states that Dendronotus iris is a
very capable swimmer and suggests that it is basically pe-
lagic, creeping on the bottom only occasionally. I suggest
that the opposite is more likely in view of the extensive
development and size of the foot as a creeping organ, and
the nudibranch’s food preference. It swims quite readily
in the laboratory or field when disturbed, and has occa-
sionally been seen to swim with no apparent provocation.
The swimming appears to be directed upwards, presum-
ably away from disturbing factors on the bottom. That
this swimming is very effective is supported by the sight-
ings of animals on the surface over 130m of water and
about half a mile from shore in Saanich Inlet.
Dendronotus robustus VERRILL, 1870
Dendronotus robustus VERRILL, 1870, Amer. Journ. Sci. Arts
50: 405 - 406; fig. 1
Dendronotus velifer Sars, 1878, Bidr. Kundsk. Norg. Arktisk.
Fauna, pp. 238 - 239, plt. 28, fig. 2
Taxonomic Remarks
There has been little doubt expressed by previous authors
(Bercu, 1894; OpHNER, 1936) that Dendronotus robust-
us is a distinct and valid species. VERRILL’s description is
complete enough to prevent confusion and for this reason
it is difficult to understand why Sars (1878) redescribed
it under a different name.
I have seen no living or preserved material of this
species, so the description is compiled from the literature,
using the two above references plus BercH (1894).
These descriptions are not very complete nor are they
specific in detail.
Body Dimensions, Texture, and Apertures
The body is stouter, less compressed laterally and less
acutely tapered posteriorly than is Dendronotus frondosus
(see plate 28, fig. 2 in Sars, 1878). The largest animal
Vol. 12; No. 4
observed was about 9cm long (Bercu, 1894), but
measured ones were 2.5cmL X 1.25cm W &X 1.25cmH
(live) (VeErRRILL, 1870) and2.8cmL x 1.0cmH 0.95
cm W (preserved) (BEeRcH, 1894). Verritt (1879,
1882) claims to have found some larger animals, but he
gives no dimensions.
The anus, atop the anal papilla, is located nearer the
second right ceras than is true for other species. The
genital openings are “as usual” (Bercu, 1894).
Apparently there is no cardiac prominence. BERcH
(1894) notes that the body is smooth, but Sars’ figures
(1878) show a scattering of small tubercles all over the
dorsal surface.
Foot
The foot is as wide as the body (VERRILL, 1870) and
small objects like hydroids can apparently be grasped
by infolding the edge of the foot.
Color
VerrILL (1870) claims the animal is a pale grayish
ground color liberally sprinkled with small yellow spots
except on the veil papillae and rhinophore stalk where
there are only a few spots. Sars (1878) and Brercu
(1894) mention a reddish body seeded with white spots.
All the lip and veil papillae are tipped with a sulphur
yellow (VERRILL, op. cit.).
Cerata
There are 6-7 pairs of relatively short, stout cerata.
They are not exceptionally arborescent, most of the
branching being only short, thick offshoots. In the first to
third pairs of cerata, the main lateral branch arises inde-
pendently of and lateral to the base of the other main
branches. In pairs 4 and 5, the lateral branch arises from
the base of the ceras, while all the posterior pairs are only
single stalks.
Rhinophore and Clavus
There is no lateral papilla on the stout, round rhinophore
stalk. The perfoliate clavus, with 10 - 12 (VeRrRILL, 1870)
or 15 - 20 (Bercu, 1894) leaves is surrounded by 5 simple,
round, smooth, tapering crown papillae of which the pos-
terior 2 are the longest.
Head, Lips, and Veil
The head is covered over by an enormous veil, upon
which there are 8-10 veil papillae. The outer two on
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Page 451
each side are the largest by about 2, but they are all
knob-like with short, simple branches on the ventral side.
About 10 small, unequally sized lip papillae adorn each
side of the lips. Some are forked, but most are simple.
Jaws
The jaw, relatively shorter than is common for Dendro-
notus, is also thicker and heavier than is usual for the
genus. No other information or diagrams have been pub-
lished.
Radula
The radula formula is 29 - 35(15 - 21:1-15- 21). Bercu
(1894) states that the median teeth of the radula are
horn yellow while the lateral teeth are colorless.
The median tooth is strongly denticulated, especially
proximally, with 15 - 20 sharp denticles that become shor-
ter towards the tip of the cusp (see plt. 2, figs. 7 - 9,
Bercy, 1894).
The hook of the lateral tooth becomes longer and more
curved from the inside toward the middle and then begins
to decrease in size as the outermost teeth are approached.
The innermost tooth (no. 1) has no denticles and virtually
no cusp. Teeth no. 2 - 4 have 1 - 2 strong denticles at the
base of the cusp. Most of the other laterals, save the outer
1-3, have 3-5 smaller denticles at the base of the
long cusp although the denticulation is irregular and often
absent. The outermost laterals are without denticles and
have only a small cusp.
Reproductive System
There is virtually no description of this system in the
literature. ODHNER (1936) mentions that the seminal re-
ceptacle is a pouch-like structure off the proximal end of
the vagina and that the prostate is a single ring of about
11 vesicles.
Geographical Distribution
So far, this species has only been found in the Arctic and
Atlantic Oceans, and has been reported from Spitzbergen,
Greenland, Siberia, northern Norway (OpHNER, 1926)
and from Cape Cod to Nova Scotia (VERRILL, 1879).
Ecology
There are no published data concerning any aspect of
ecology except that Dendronous robustus occurs from the
intertidal down to at least 200m (OpHNER, 1939).
Page 452
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Vol. 12; No. 4
Dendronotus dalli BErcH, 1879
(Plate 63, Figure 31; Text figures 1, 4 - 6, 13 - 15)
Dendronotus dalli BERGH, 1879, Proc. Acad. Nat. Sci. Phila-
delphia 5(1): 150; plt. 1, fig. 21; plt. 2, figs. 9 - 12;
plt. 3, figs. 2-6
Dendronotus elegans VERRILL, 1880, Proc. U.S. Nat. Mus. 3:
385 - 386
Dendronotus frondosus (Ascanius, 1774). OpHNER, 1936,
Mém. Musée Roy. Hist. Nat. Belg., ser. 2, 12 (3):
1105 - 1109
Taxonomic Remarks
Bercy (1879) first described Dendronotus dalli from a
“bulbus pharyngeus’”’ or buccal mass dredged in the Bering
Strait. The main points of difference from D. frondosus,
he felt, were the complete lack of denticulation on the
median tooth and the more distinct denticles on the masti-
catory border of the jaw.
In a later paper that included a more extensive descrip-
tion of a whole animal, BercH (1894) expressed some
doubt about the validity of Dendronotus dalli. He indi-
cated that it, like D. purpureus, may be a color variety of
D. frondosus that lacks all the median tooth denticulation.
However, he retained the name D. dalli in a still later
description of a specimen collected at Bear Island (BERGH,
1900).
O’DonocHvE (1921) gave a brief description of the
external characters of a small specimen. However, this
description, with the exception of the larger number of
lateral teeth, fits Dendronotus spec. nov. almost exactly.
O’DonocHUE deposited the specimens, upon which he
based his description, in the museum at the Nanaimo
Biological Station, but during a subsequent move, they
were apparently lost (Clarke, pers. comm.; Quayle, pers.
comm.).
OpHNER (1936) expressed some doubt about O’Dono-
GHUE'’s description and concluded that, without more spe-
cific and detailed information, Dendronotus dalli should be
considered a synonym of D. frondosus. He dismissed all
of Bercu’s work by stating that the radula characters
upon which Bereu based his descriptions are too variable
for taxonomic purposes (ODHNER, 1926, 1936). MacFar-
LAND (1966, p. 257), in agreeing with OpHNER, stated:
“He [O’DonocHUvE] gives no anatomical details to support his
conclusion [that Dendronotus dalli is a distinct species], and
until these are produced, the conclusion of OpHNER (1936)
that the radula characteristics are too variable to justify spe-
cies discrimination are most reasonable.”
VERRILL’s (1880) description of Dendronotus elegans
strongly suggests that the animal was a D. dalli and not
a morphological variant of D. frondosus as ODHNER
(1936) suggests. Because there are no further records pub-
lished by VERRILL, I suggest, on the basis of this descrip-
tion, that D. elegans be considered a synonym of D. dalli.
The following description shows that Dendronotus dalli
is indeed a valid species, distinct morphologically and bio-
logically from D. frondosus.
Body Dimensions, Texture, and Apertures
In the relatively heavy, laterally compressed, limaciform
body, the rounded dorsum merges indistinctly with the
vertical sides. The posterior third of the body tapers to
form a short, moderately blunt-pointed tail (Figure 1).
The largest specimen was 13.5 cm L and there were 11
others over 9.0cm L. Most of the animals were 4.0 - 6.0
(1.9- 13.5) em L X 0.7- 1.5 (0.6 - 3.1) cmH x 0.5 - 1.0
(0.4 - 1.6) cm W. A few representative measurements of
live animals were: 4.4cm L < 0.7cm H X 0.5cm W;;
65cm L & 1.2cmH X 0.9cm W; 8.0cm L & 1.5cm
H x 1.2cm W. Bercu (1879, 1894) estimates one ani-
mal to be 10 cm L while others were 3.5 cm and 4.2 cm
L, both preserved. O’DonocHuE’s (1921) largest speci-
men, preserved, was 3.1cm L & 0.9cm H X 0.6cm W.
VoLopcHENKO (1955) reports specimens up to 10 cm L.
The anal papilla, if present, is inconspicuously located
halfway beween the bases of no. 1 and no. 2 right cerata.
The conspicuous genital openings are located about
halfway up the right side of the body, just anterior to the
first ceras.
The cardiac prominence is usually only a slight eleva-
tion, but in a few specimens it is protruded up to 4 - 5 mm
above the dorsum. Only 2 of 60 animals had any body
papillae and these were limited to the cardiac prominence.
All the other animals were smooth.
Explanation of Plate 63
Figure 29: Dendronotus frondosus (AscANius), white form from
20 m off San Juan Island, Washington X 2
Figure 30: Dendronotus iris Cooper, grey form. In this particular
animal, the grey is overlaid with the muddy brown pigment com-
monly seen on animals from deep-water muddy bottoms x 4
Figure 31: Dendronotus dalli Brercu. Note the relatively short,
stout, but extensively branched cerata (CE) and veil papillae
(VP) x1
Figure 32: Dendronotus rufus O’DonocHuE. Note the tall, ex-
tensively branched cerata (CE) and rhinophores (RH). The magenta
pigment is less dense on the veil papillae (VP) and rhinophores X 4
THE VELIGER, Vol. 12, No. 4 [RopILLiARp] Plate 63
Figure 29
Figure 32
Figure 31
Figure 30
me
ir
Vol. 12; No. 4
Foot
The foot (Figure 1) is about 4 as wide as long. The
anterior end is blunt and rounded while the posterior
end tapers abruptly to a blunt point. When the animal is
crawling, the foot flares laterally along its whole length
though not as extensively as in Dendronotus iris. During
swimming, the edges fold quite tightly together. The sole
of the foot is white or very pale pink with a faint hint
of darker pink around the edges; this edging is not as
prominent as it is in D. iris or D. rufus.
Despite the large size of the foot, the animal is capable
of clinging stubbornly to the stiff branches of the hydroids,
Abietinaria rigida or A. amphora. It is able to maintain
its position on plain rock in the face of a relatively
fast current.
Color
Previously, all the descriptions have been of preserved
specimens which become a dull, indifferent gray in alcohol
or formalin. In life, Dendronotus dalli is rather uniformly
colored in varying shades of translucent white through
pearl pink to salmon pink; 3 specimens were a mauve
hue. Most of the animals collected were pearl pink (Plate
63, Figure 31). The color is lightest on the sides, becoming
darker on the dorsum and darkest on the cerata, rhino-
phores, and veil papillae. The extremities of these proces-
ses are usually covered with an opaque, dead-white pig-
ment, occasionally extending down the main branches in
irregular lines of scattered patches. A few animals were
sparsely to liberally sprinkled with tiny, opaque, white
spots.
The clavus and oral region (outer lips and lip papillae)
are a yellowish color. The tan to dark pink liver and the
white gonad are plainly visible through the body wall.
Cerata
The short, but extensively branched cerata are in 6-7
(4-8) pairs arranged at intervals along the dorso-lateral
margin of the body. VoLopcHENKo (1955) reports up to
9 pairs. Both the interval and size of the cerata gradually
decrease from anterior to posterior. On the tail, there are
1 - 2 (1-5) unpaired, but often relatively large, branched
processes or “accessory cerata.” These occasionally occur
between the other cerata also.
The main branches are quite stout, but short; often the
total height of the cerata is not much more than the
height of the body. The main branches give rise to even
shorter, stout secondary branches which in turn may
divide into small tertiary branches. These secondary and
tertiary branches, because they are short and arise from
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Page 453
all sides of the main branches, impart a “fuzzy” appear-
ance to the cerata. When viewed from above, the ceratal
mat provides very little cover for the dorsum.
The branching pattern is “fan-shaped” (Figure 4a)
with 4 (3-4) main branches in no. 1 cerata, 3 - 4 in no.
2, 3 (2-4) in no. 3, 3 (2-3) in no. 4, 2-3 in no. 5, 2
(1-2) in no. 6, and 1 in nos. 7 and 8. Where there are
4 main branches, there is a dichotomy at or very near the
base of the ceras. Each of these stalks divides almost im-
mediately to form 4 branches. The medial branch is short,
the next one out (no. 2) is the longest, no. 3 is about
equal to no. 2 and the lateral branch (no. 4) is shortest,
being about 4 to 4 as long as no. 3. Where there are 3
main branches, the lateral branch (no. 3) arises at or near
the base of the ceras above which the ceras bifurcates to
form branches no. 1 and no. 2. The medial branch is the
longest, but no. 2 is only slightly shorter while the lateral
is again about 4 to 4 as long as no. 1. If there are only 2
main branches, they divide at the base and no. 2 is vari-
able in length. In some large specimens, there is an extra
medial branch in the first pair of cerata.
The cerata of Dendronotus dalli appear to be subject
to many aberrations in size and branching. This may be
the result of attacks by predators, probably fish.
Rhinophore and Clavus
The rhinophore, like the cerata, is relatively short and
stout with many small branches.
The lateral papilla, arising from or near the base of
the rhinophore stalk, is heavy and long, often reaching
higher than the extended clavus. The degree of branching
varies a good deal from a few small, simple papillae
arising from the main branch to the extensive secondary
and tertiary branching comparable to the cerata.
The yellow, perfoliate clavus has 20-24 (16-33)
roughly alternating, shallow-cut leaves. The top 3 - 4 and
bottom 2 - 3 leaves are small and equal. The large leaves
almost completely encircle the clavus while the small ones
are limited to the posterior border.
Extending upward from the rhinophore sheath are 5
(4-12) highly variable crown papillae. They are usually
long, but not particularly stout. All are about equal in
length though the posterior two may be 2 longer than
the others, and often as tall as the rhinophore stalk itself.
The branching consists of a few relatively short papillae
arising from the proximal half of the main stalk.
Head, Lips, and Veil
The veil, not sharply marked off from the head, bears 4
or, more often, 5 pairs of stout, extensively branched veil
Page 454
papillae radiating fan-like from it (Figure 1). The ar-
rangement and size of the veil papillae varies, making it
difficult to count and describe them. In those animals
with 4 pairs of papillae, the medial pair (no. 1) is usually
the longest, no. 2 is about 4 to 3 as long as no. 1, no. 3 is
about equal to no. 1, and no. 4 is the shortest at a } to 4
the length of no. 1. When 5 pairs are present, no. 1 is the
longest. No. 2 pair may be + to ? as long and originates
ventro-lateral to no. 1. Or, it may be about equal to no. 3
in size. Pairs no. 3 and no. 4 are usually about 4 to ¢ as
long as no. 1. The unbranched lateral pair, no. 5, is about
4 to tas long as no. 1. In a few specimens all 4 or 5 pairs
of veil papillae are approximately the same length.
Many (15-30+) simple lip papillae, ranging in
length from mere tubercles near the mouth to long, finger-
like papillae on the lateral margins, adorn the oral surface
(Figure 1). Some of the larger lip papillae, especially
those just ventral to the veil papillae, may be forked at
the tip or, rarely, they may be quite bushy. All the lip
papillae are sensitive and able to retract when disturbed.
Jaws
The jaws of Dendronotus dalli (Figure 13) seem longer
and stronger than they are. The buccal mass has a
length : width ratio of 1.2 - 1.4 and a length : height ratio
ofp Ibo) ae 7:
The jaws are transparent, citron yellow, slightly darker
on the hinge and masticatory process. In his original de-
1.0mm D
Figure 13
Dendronotus dalli jaw
B. Ventral view C. Inside lateral view
D. Outside lateral view
A. Dorsal view
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Vol. 12; No. 4
scription, BercH (1879) incorrectly states that the jaw is
darker than that of Dendronotus frondosus, but in sub-
sequent papers (1894, 1900), he indirectly corrects this
error.
The body has a deeply convex, mytiloid form, widest
in the posterior third and bluntly convex on the posterior
end. The length : width ratio is about 2: 1 (Figure 5 for
measurements). The large, strong, deeply grooved dorsal
process, about 0.45 the jaw body length, is inclined at
a 60° angle to the longitudinal axis of the jaw body.
The process is joined to the body by a strongly convex ex-
pansion for less than half of its length. The short masti-
catory process, only slightly arched, bears about 200 low
denticles (BrRcH, 1894) and is joined to the body by a
thick, slightly concave expansion.
Figure 14
Dendronotus dalli radula
Row 21 in a radula with 38 rows of teeth
Radula
The radula (Figure 14), ODHNER’s (1926, 1936) com-
ments notwithstanding, is distinctly different from that of
Dendronotus frondosus. When the figures from all his
descriptions are compiled, BercH (1879, 1894, 1900)
reported a radula formula of 42 -51(11-16-1-11- 16)
for D. dali. O.DoNocHUuE (1921) found it to be 38 - 40
(10-14-1-10-14). From a study of 29 radulae, I ob-
tained a radula formula of 37-48 (9-14-1-9-14). The
composite formula is 37-51 (9-16°1-9-16).
Bercu (1894) stated that the median teeth are a dark
mahogany color but I found only clear or very slightly
yellow teeth in fresh specimens. f
The cusp of the median tooth, rounded at the apex
and weakly concave on the sides, becomes taller towards
the radula sheath. In the oldest teeth the width : height
ratio of the cusp is about 2:1, whereas from about row no.
20 on it is 1.7- 1.9: 1 (Figure 6 for measurements). As
maintained by Bercu (1879, 1894) and O’DonoGHUE
(1921), there are no conspicuous denticles on the median
teeth. Except for a few cases, there is only an indistinct,
Vol. 12; No. 4
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Page 455
irregular serrulation at most and this disappears toward
the apex. A few specimens had 30-40 tiny, distinct,
regularly arranged denticles on each side of the cusp,
usually in the posterior half of the radula.
The lateral teeth are long, narrow, flat plates increasing
in size from anterior to posterior. The cusps of the lateral
teeth are slightly inclined (about 10°) toward the midline
and away from the long axis of the tooth’s base. On the
innermost laterals, the cusp is about equal in length to the
denticles. The next 2 - 4 teeth have large, sharp, weakly
curved cusps bearing denticles along the proximal, lateral
border. The sharp, thin cusps on the outer teeth decrease
in length and finally disappear from the outer 1 - 2 teeth.
Of the 9 - 14 lateral teeth in the specimens I examined,
only the outer 2-3 (1-5) were without denticles. All
the rest have 3 - 7 (1 - 19) sharp, irregularly arranged and
shaped denticles that are long and heavy when there are
only a few (1-6), but more slender when there are many
(7- 11+). There is a tendency for the number of den-
ticles to decrease from the inner to the outer teeth. On the
inner two laterals (no. 1 and no. 2), there are respectively,
4-5 (2-11) and 4-6 (2-10) denticles. There are 4-6
(3-11) denticles on no. 3, 3-6 (2-10) on no. 4, 3-5
(2-8) onno. 5, 2-4 (1-8) onno. 6, 1-4 onno. 7, 1-3
on no. 8 and no. 9, and sometimes 1 or 2 small ones on
no. 10. Numbers 8 to 10 occasionally lack any denticles.
Reproductive System
In Figure 15, the reproductive system is spread out to
show the relative size and relationships of the component
parts.
Each lobule of the hermaphrodite gland seems to have
a dark gray patch on the distal end. The hermaphrodite
duct links the gland to the relatively long, narrow, coiled
ampulla which merges indistinctly into the short, wide
spermoviduct. The short proximal portion of the vas de-
ferens expands into the prostate consisting of many small,
usually spherical alveoli arranged as a flattened sphere
concentric with the vas deferens. The wide, quite short
and muscular distal portion of the vas deferens tapers
very slightly before entering the penis. The penis is bullet-
shaped, thick at the base and tapered to a point in the
distal third.
The short, wide vagina narrows slightly before entering
the vestibule relatively far from the common female gen-
ital opening. The small bursa copulatrix opens into the
vagina near its distal opening. The seminal receptacle,
a flattened ovoid structure that is bent back on itself,
is located at the proximal end of the vagina. The insemi-
nation duct appears to enter the vagina about halfway
between the bursa copulatrix and the seminal receptacle,
1.0mm
|
HD
Figure 15
Reproductive system of Dendronotus dalli (exploded view)
AMP = ampulla BC = bursa copulatrix
DVD = distal vas deferens ED = ejaculatory duct
FC = (to) fertilization chamber FGA = female genital aperture
FGM = (to) female gland mass HD = hermaphrodite duct
ID = insemination duct OV = oviduct P = preputium
PE = penis PR = prostate PVD = proximal vas deferens
SO = spermoviduct SR = seminal receptacle VA = vagina
VE = vestibule
but closer inspection reveals that it continues as a tube
within the lumen of the vagina and actually opens near
the mouth of the seminal receptacle. The other end of the
insemination duct is confluent with the short, transparent
oviduct forming a fertilization chamber.
Geographical Distribution
Except for one specimen, this species has only been col-
lected in the northeastern Pacific and the Bering Straits
Page 456
at depths of 10 - 60 fathoms. Bercu (1879, 1894, 1900)
had specimens from Bear Island in the Barents Sea (1),
the North Pacific (2), and Bering Strait (1). O’Donoc-
HUE (1921) collected them at Halibut Bank, Nanoose
Bay, and Gabriola Pass, all off eastern Vancouver Island,
B. C. VotopcHENKo (1955) mentioned them as occur-
ring in the North Pacific off the Russian Coast.
Besides the Victoria Breakwater (50 animals), I col-
lected them from Albert Head, Victoria (3), Eagle Point
(6), Lonesome Cove (5), Harney Channel (4 dredged),
Turn Rock (3), and San Juan Channel (4). The latter
5 areas are in the San Juan Archipelago (see Appendix I).
Ecology
Up to now, nothing has been published concerning the
ecology of this species.
Adults, obtained from June, 1966 to April, 1967, were
largest and most common from October to November,
1966 (although this may be a reflection of the collection
pressure and timing). They were all collected at 20 - 30m
from rocky areas supporting large populations of Abietin-
aria rigida and A. amphora, but few other hydroids
(with the exception of Lonesome Cove). These areas,
often bordered by sandy or muddy bottoms, are usually
swept by moderate currents. As mentioned above, Dend-
ronotus dalli is able to cling to the hydroids despite these
currents.
During a dive off the Victoria Breakwater on August
10, 1966, large numbers of Dendronotus dalli egg masses
were seen. White, loosely coiled, they were deposited on
Abietinaria rigida, the rocks, and on other sessile animals
such as Balanus nubilis and the bryozoan, Heteropora sp.
During a second dive in late October, 1966, only a few
egg masses were seen and in December, 1966 and January,
1968, there were none. One egg mass was collected from
Eagle Point on March 27, 1967 and another from Turn
Rock on January 21, 1968.
The animals collected in December, 1966, when put in
the water tables at the Friday Harbor Laboratories, began
to copulate and deposit eggs all during January, 1967. It
is difficult to tell whether this was a response to the change
in light, temperature, salinity, or substratum; in any case,
the gonads were mature when they were collected. A
sample from some of these white to cream-colored egg
masses contained 4-6, 2-3, 2-3, 3-4, 1-3, and 6-9
eggs per capsule. Hatching, in 4 cases, took 13-17, 15-18,
17 - 18, 18 - 19 days.
To the human diver’s eye, these nudibranchs are visible
up to 20 feet against the dark background of the rock and
hydroid bed. Presumably fish are also able to see them
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Vol. 12; No. 4
and, judging from the number of animals with mutilated
cerata and rhinophores, probably attack them rather
frequently.
Dendronotus dalli is capable of effective, if not pro-
longed swimming when it is disturbed. So far, only me-
chanical stimuli such as pinching the cerata has proved
effective in eliciting the response. None of the potential
predators such as starfish, other carnivorous gastropods,
or crabs have shown any effect although, in nature, the re-
action to pinching seen in the laboratory may be an effec-
tive escape response from a crab. No other methods of de-
fense were obvious, but epidermal secretions are a possi-
bility (EpMunps, 1966; THompson, 1960b).
Dendronotus rufus O’DONOGHUE, 1921
(Plate 63, Figure 32; Text figures 4 - 6, 16 - 18)
Dendronotus rufus O’DoNocHuE, 1921, Trans. Roy. Canad.
Inst. 13 (1): 190 - 192; plt. 3, figs. 25 - 27; plt. 4, fig. 48
Dendronotus frondosus (Ascanius, 1774). OpHNER, 1936,
Mém. Musée Roy. Hist. Nat. Belg., ser. 2, 12 (3):
1105 - 1109
Taxonomic Remarks
O’DonocHUE, in describing this species, makes only brief
and uncritical reference to the important taxonomic char-
acters. It is little wonder that ODHNER (1936, p. 1108)
said:
“D. rufus ... has a colour which makes it a probable syn-
onyme [sic] of D. frondosus, from which it cannot be separated
(according to the description) in external characteristics.”
Probably because of his belief that the radula characters
in this genus are too variable to be of any use taxo-
nomically, ODHNER says nothing about the differences,
pointed out by O’DonocHuE, between the radula of
Dendronotus rufus and D. frondosus.
O’DonocHueE’s (1921) largest specimen was only 1.4
cm L while my smallest one was 7.0 cm L and the average
was 17cm L. It is this size discrepancy that made it
difficult to determine if the species described below is the
same as that described by O’DonocHuE. The presence
of a red line around the foot and the color of some of
his specimens are the most important characters that
link my specimens to Dendronotus rufus.
The type specimen has apparently been lost (Quayle,
pers. comm.; Clarke, pers. comm.; Tebble, pers. comm.),
probably during a move of material from the Nanaimo
Biological Station to the Canadian National Museum at
Ottawa, Canada.
Vol. 12; No. 4
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Page 457
Body Dimensions, Texture, and Apertures
The limaciform body is relatively “heavy,” but not as
stout in appearance as that of Dendronotus iris despite
the generally larger size of D. rufus. The high, rounded
dorsum is confluent with the vertical sides with no sign
of a border except the cerata. The animal is laterally
compressed, though not markedly so, and tapers to a
short, blunt tail.
As noted above, O’DoNocGHUE’s specimens were small,
the largest being 14cm L & 0.45cm H X 04cm W
and the smallest 0.6cm L. He subsequently collected a
“fairly large” specimen, 1.65cm L & 0.52cm H X 0.36
cm W (O’DonocHuE, 1922, p. 164). The largest animal
of 26 measured in this study was 28.0cm L & 4.5cm H
< 4.2 cm W while the smallest was 7.0cm L & 1.5cm H
x 1.0 cm W. Most of the animals were 15.0 - 20.7 (7.0 to
28.0) cm L X 3.0-4.0 (1.2-4.5) cmH X 2.5 - 3.0 (1.0
to 4.3) cm W.
The anus is on a distinct papilla located about halfway
between the first and second right cerata. The top of the
papilla is usually magenta like the cerata.
The genital apertures, located about halfway up the
side and just anterior to the first ceras on the right, may
be quite distinct, depending upon the sexual condition of
the animal.
The cardiac prominence is usually indistinct. The body
may be smooth or it may have a few small, magenta-
tipped, sub-conical papillae widely scattered on the dor-
sum and the sides. Occasionally, it takes on a wrinkled
appearance, particularly if the animal is at all contracted.
Foot
The foot, about 6 longer than wide, is bluntly rounded
anteriorly, while the posterior end is bluntly pointed. The
edge flares slightly when the animal is crawling on a
smooth surface.
The sole of the foot is white. Around the dorsal edge of
the foot is a distinct light to dark magenta line made up of
small pigment patches. In a few light-colored specimens,
this line was not outstanding but it was discernible. In
the 2 brick-red animals, this line was dark pink, but
because of the body color, it was not so obvious.
Observations in the field and in the laboratory indicate
that this species is only partly successful in maintaining
its position in a moderately strong current. It also appears
unable to grasp the thin stalks of a hydroid, although,
considering the size of the animal, this seems reasonable.
As yet, nothing is known about the function of the foot
in the juvenile stages.
Color
Because color is the important link between O’Donoc-
HUE's description and mine, his original account is quoted
in its entirety (O’DonocHuE, 1921, p. 190):
“In the living animal, there is a considerable range of colour
variation. The general body colour varies from a transparent
grey through a semi-transparent pink to a deep brick red. In
the grey form, the cerata were of a pale pink colour, in the
pink form there was a narrow line of darker, more opaque
pink, running around the edge of the foot and up around the
head at the anterior end. A similar, but less marked darker
line was also present in the red form.”
All except 2 specimens that I examined had an opaque,
gray-white ground color but, in a few, a liberal sprinkling
of very fine magenta dots and some small, magenta-tipped
papillae imparted a pinkish hue to the animal (Plate 63,
Figure 32). Except in these few extensively pigmented
animals, the gray-white color extends for varying distan-
ces up the cerata, rhinophores, veil and lip papillae. It
merges with a deep magenta pigment which becomes
darker and more concentrated towards the distal ends of
the above processes. In some animals, the magenta color
becomes almost black. In others, it is washed out giving
the appendages a distinctly yellow tinge under the ma-
genta. The magenta line around the foot has been men-
tioned above.
Two specimens were of the “red form” mentioned by
O’DonocuHvE (1921). The ground color is a deep brick-
red, lightest on the sides, darker on the dorsum and bases
of the appendages, and darkest on the distal portions of
the appendages. Scattered over the body and appendages
are many small, opaque, yellow to white spots. A few
larger, pale cream patches were present on the sides just
below the bases of the cerata. Between the cerata, there is
an irregularly blotched line of large, opaque white pat-
ches extending from the rhinophore to the tip of the tail.
Cerata
The cerata of this species are by far the largest and most
dendritic of any of the species of Dendronotus. In an ani-
mal 14.0 cm L, the tallest cerata were up to 8cm tall
while the widest part of the “ceratal mat” was over 8 cm
across. When viewed from above, the body of the animal
is almost completely covered by the cerata.
There are 7 - 8 (6-9) pairs of cerata placed at slightly
decreasing intervals and followed by a single, unpaired
ceras in a few animals. The cerata decrease in size also,
the last pair being about 4 or less as tall as the first.
Between the main cerata are found 4-11 “accessory
cerata” varying in size from short, simple tufts to arbor-
Page 458
escent branches nearly as large as the smaller cerata, and
pigmented much like the cerata. Although most of them
are irregularly placed, there are usually 2-4 in a trans-
verse row across the cardiac prominence.
The main branches of the cerata are thick, heavy, and
very long. Originating from them are many long and
relatively stout secondary branches which in turn divide
into long, thin tertiary branches, or which terminate in
a bushy tuft of fine filaments. The tertiary branches, fine
as they are, often divide to form a smaller bushy tuft.
The pattern of branching is “fan-shaped” (Figure 4a).
There are 4 main branches in the first and second pairs
of cerata, 3 - 4 in the third, 3 in the fourth, 3 (1-3) in
the fifth, 2 (1 - 2) in the sixth, and 1 (1 - 2) in the seventh
and eighth pairs. The main branches all split off the very
short stalk at about the same level. The medial branch
(no. 1) and no. 2 are about equal in size, while no. 3 is
about # as large and no. 4 is about $ to 2 as large as no. 1.
When 3 branches are present, the proportions are about
the same but there is only one long medial instead of 2.
Because of the great length of the secondary branches,
it is very difficult to determine what to count as main
branches and the choice becomes arbitrary especially in
the anterior pairs. There is usually a large medial second-
ary branch arising from the base of main branch no. 1
in the anterior 1 - 4 pairs of cerata. This and a smaller,
lateral, secondary branch of main branch no. 4 (or no. 3)
of the anterior pair(s) of cerata are particularly difficult
to separate from the main branches.
The hepatic diverticula extend into the rhinophores as
well as the anterior 4 - 5 pairs of cerata which lie dorsal
to the liver. These are visible only after dissection and are
limited to the medial 2 - 3 branches of the ceras.
O’DonocHuE (1921) described 5 pairs of cerata. The
first 3 pairs had 3 simply branched trunks, the fourth pair
had 2 simple main trunks, and the members of the fifth
pair were simple projections.
Rhinophore and Clavus
The rhinophore, like the cerata, is extensively branched
and very tall, reaching as high as the first pair of cerata.
The heavy, firm lateral papilla originates from the base
of the stout rhinophore stalk and, despite being at a 30°
angle to the stalk, reaches as high as the clavus or even
the top of the shorter crown papillae. From the main
stalk spring many long secondary branches which may
end in bushy tufts or may give rise to slender, tertiary
branches that finally end in tufts.
The sub-conical, yellowish to purple-spotted clavus is
quite small relative to the size of the animal. There are
19 - 24 alternating shallow-cut leaves as opposed to the
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Vol. 12; No. 4
8 - 10 non-alternating leaves mentioned by O’DoNoGHUE
(1921). The upper 2 - 4 and lower 4 - 5 leaves are small
and equal while on the rest of the clavus the large leaves
are about 2 wider than the smaller, and almost com-
pletely encircle the clavus.
Radiating from the top of the sheath, like wheel spokes,
the 5 crown papillae are long and stout. The branching,
usually from the distal half of the papillae, may be re-
stricted to a few relatively simple papillae and tufts or it
may be more extensive albeit less so than on the other
appendages. The crown papillae are all about equal in
length; the longest, the postero-medial, is rarely more
than 2 longer than the shortest. Occasionally, the 3
posterior ones are about 14 longer than the anterior 2.
Head, Lips, and Veil
The 5 pairs of veil papillae are made up of very dendritic,
long, thick branches that radiate in an antero-dorsal
plane at about 30° from the horizontal. The main
branches divide into numerous secondary and tertiary
branches just as already described for the cerata and
lateral papillae. This type of branching makes it exceed-
ingly difficult to determine what is and what is not a veil
papilla. To make it even more difficult, the thick bases
of the veil papillae arise immediately adjacent to each
other. There is not even a gap in the midline between veil
papillae from two sides as there is in the other species.
The medial pair of veil papillae (no. 1) is the largest
with no, 3 only slightly shorter. Number 2 pair, originating
just ventral and lateral to the base of no. 1, is about } to
4 as long as no. 1. Pairs no. 4 and no. 5 are about equal to
no. 2, though no. 5 is usually somewhat shorter.
Immediately ventral to the veil papillae are 5-10
large, branched papillae that I have called “accessory
lip papillae.” This name may seem incongruous consider-
ing that these papillae are far larger and more branched
than the other lip papillae, but they appear to function
as veil, rather than lip, papillae; hence the term “acces-
sory.”
Posterior to these accessory lip papillae are the true lip
papillae. There are 30 or more of these short to long,
finger-like projections. A few of the anterior ones may be
forked or slightly branched on top. Arranged in vaguely
parallel rows coursing fore and aft on either side of the
lips, they are longest laterally and shortest near the
mouth.
Jaws
The jaws (Figure 16) and buccal mass of Dendronotus
rufus are remarkably small for so large an animal. From
Vol. 12; No. 4
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Page 459
a sample of 8 buccal masses, the length of the buccal mas
was found to be about 1/20 to 1/25 the total length of
the body.
1.0mm D
Figure 16
Dendronotus rufus jaw
B. Ventral view C. Inside lateral view
D. Outside lateral view
A. Dorsal view
The light yellow jaws are relatively thin and delicate.
The body is a moderately convex, mytiloid form, slightly
wider in the posterior third and about 2.2 longer than
wide. The dorsal process, about 0.45 < as long as the body
and inclined at a 70° angle with its long axis, is joined
to the body by a strongly convex expansion throughout
half or more of its length. The distal portion of this broad,
heavy dorsal process is curved downward and the dorsal
groove is deeper and wider than in any other species.
The masticatory process, at a 75° angle with the long
axis of the body, is joined to the body by a strong, shal-
lowly concave expansion and bears numerous rows of
narrow, low denticles.
Radula
O’Donocnur (1921) described a radula formula of
32-35 (6-8-1-6-8): the laterals had 3-6 stout den-
ticles while the medians bore 10 large sharp denticles. In
contradistinction to this, in the 9 radulae that I examined,
the formula was 33 - 35 (10 - 16:1-10- 16). The laterals
have a variable number of short denticles, and the medi-
ans bear many short, blunt denticles (Figure 17). If the
animals O’DonoGHUE describes are juveniles of this spe-
cies, then it seems that there must be a marked change in
the radula characteristics during an individual’s life; it
may be these transition stages which cause the confusion
about which OpHNER (1926) writes.
The cusp of the median tooth, rounded or bluntly
pointed at the apex and very weakly bulged on the sides,
is usually about 1.5 - 1.7 as wide as long, but in a few
posterior teeth the ratio may increase to 2 (Figure 6
for measurements). The denticulation of the cusp varies
considerably. There are commonly 16 - 22 (6-28) small,
blunt, more or less regularly spaced, denticles diminishing
in size towards the apex and sometimes disappearing al-
together. The indentations of the proximal denticles con-
tinue on the dorsal surface of the cusp as fine, shallow
furrows, giving the tooth a delicately corrugated appear-
ance. In a few specimens, there is only a fine serrulation
along the edge while in a few others, the older teeth may
be quite jagged where pieces have apparently been broken
off. On any particular radula, the number of denticles on
the median tooth generally increases from the oldest teeth
to the newest.
0.1 mm
—
Figure 17
Dendronotus rufus radula
Rows 21 and 22 in a radula with 36 rows of teeth
Most of the tall, stout lateral teeth bear a strong, blunt-
ly-pointed, straight cusp markedly inclined (20 - 30°)
toward the midline. The inner few teeth have a distinct
protuberance on the lateral border at the base of the cusp.
In a transverse row, the cusp increases in size from a short,
often insignificant quantity in teeth no. 1 and no. 2 to a
maximum length in teeth no. 6 to 8 and then decreases
again, often disappearing in the outermost | or 2 teeth.
The denticulation of the lateral teeth is quite variable.
Generally, the short, sharp denticles do not project very
far above the lateral border of the cusp and then continue
down the face of the base as relatively deep, distinct
furrows. There is a vague tendency for the number of
denticles per tooth to increase from anterior to posterior
rows while within a row it is greatest at teeth nos. 4 - 7,
and decreases on either side. The following list of denticle
Page 460
numbers per lateral tooth is compiled from counts made
on 6 rows at various points along the radula for 9 radulae.
The numbers in parentheses represent extremes while
those outside represent 70% of the teeth.
Lateral Tooth (numbered Number of Denticles
from medial to lateral)
1 none 50% of time;
1 - 4 otherwise
2 0-3 (0-5)
3 0-3 (0-6)
4 2-4 (0-6)
5 2-5 (0-7)
6 1-6 (0-8)
7 3-5 (0-9)
8 0-6 (0-9)
9 0-3 (0-8)
10 0-2 (0-7)
11 0-1 (0-6) .
12 0-1 (0-2)
13 0
14 0
15 0
Reproductive System
The reproductive system (Figure 18) may be concisely
described as “long.”
The large, white hermaphrodite gland of sexually ma-
ture animals appears gray in immature animals because
of the small black spots on the lobules. The hermaphrodite
duct expands suddenly to form a long, narrow ampulla,
exceeded in relative length only by that of Dendronotus
iris. The ampulla narrows rapidly to form the short sperm-
oviduct that bifurcates to form a very short, transparent,
rugose oviduct and an equally short proximal portion of
the vas deferens.
The vas deferens expands to form the prostate, a rela-
tively enormous sphere concentric with the vas deferens
and comprised of many small to large irregularly shaped
alveoli. From this large prostate arises a very long, nar-
row, extremely convoluted and coiled distal portion of
the vas deferens. It tapers very gently throughout its whole
length to become a thin tube. As such, it enters the base
of the penis to form the long, convoluted ejaculatory
duct. The long slender penis, coiled and folded back on
itself in the preputium, is tapered to a slightly bulbous
tip. The male portion of the Dendronotus rufus genital
apparatus is most distinctive: the prostate is larger, the
vas deferens is longer and more convoluted, and the
penis is relatively longer and thinner than in any other
species in the genus.
THE VELIGER
Vol. 12; No. 4
Figure 18
Reproductive system of Dendronotus rufus (exploded view)
AMP = ampulla BC = bursa copulatrix
DVD = distal vas deferens FC = (to) fertilization chamber
FGA = female genital aperture FGM = (to) female gland mass
HD = hermaphrodite duct ID = insemination duct
OV = oviduct P = preputium PE = penis
PR = prostate PVD = proximal vas deferens
SO = spermoviduct SR = seminal receptacle VA =vagina
VE = vestibule
The long but narrow and tortuous vagina terminates
proximally in a large, nearly sessile, ovoid seminal recept-
acle. The bursa copulatrix, not much wider than its long
stalk, is located near the opening of the vagina into the
vestibule. A very short insemination duct is applied to the
outer surface of the vagina for a short distance before
joining the oviduct.
Geographical Distribution
O’DonocHuE (1921, 1922, 1924) collected Dendrono-
tus rufus at Departure Bay, Nanoose Bay, Breadwater
Island, and Thetis Island, all in the Strait of Georgia, B.
C. The depth range was 20 - 40m.
Vol. 12; No. 4
THE VELIGER
The specimens collected for this study were obtained
at Alki Point, Seattle (23 animals) from 15 - 17m; De-
parture Bay, Nanaimo (1 animal) from 20m; Brown
Island in Friday Harbor (1 animal) from 7m; and the
floats at Jensen’s Boat Yard, Friday Harbor (2 animals).
Ecology
Juveniles of this species have not been seen, but adults
were found from November, 1966 to March, 1967 and
again from January to March, 1968. Dendronotus rufus
seems to prefer areas of slight to moderate current action
possibly because of the apparent inability to hold on in
swift currents. The substratum has usually been rock sur-
rounded on all sides by sand or mud or both.
Examination of the fecal pellets plus observations in
the field and laboratory indicate that scyphistomae, when
present, form part of the diet of the Alki Point popula-
tion of Dendronotus rufus. Nothing else was found in their
guts except for a few amphipods that occurred on the
scyphistomae. The fecal pellets of the animals collected
at areas other than Alki Point contained no nematocysts
or anything else recognizable.
During late January, 1967, nudibranchs collected two
months previously began depositing egg masses in the
aquarium. Shortly after, on February 2, 7 egg masses were
seen at Alki Point on the rocks and pipe. The animals
collected then plus the previous group (16 in all) laid
about 60 nidosomes up to the end of March when they all
died. During January and February, 1968, about 100
copulating pairs were seen at Alki Point and 175 - 200
egg masses were deposited during February. By March 20,
1968, all the animals plus the egg masses were gone. In
the field as well as the laboratory, the white to cream
colored egg string was deposited in a regular though loose
spiral. The number of eggs per capsule varied from 4 - 5
to 17 - 27, with an average of 13 - 17. The length of time
from oviposition to hatching of the veligers varied from
13 - 20 days with most hatching in 15 - 18 days.
Only because of its large size is Dendronotus rufus easy
for a diver to see in the field. The dark cover provided
by the cerata helps the animal to blend in with the back-
ground, at least those upon which it has been seen.
Dendronotus rufus appears to be a very poor swimmer
responding quite slowly to strong mechanical stimulation.
The species has a possible defense mechanism that the rest
of the Dendronotus species lack: a thick, extremely sticky
mucus. This mucus makes the animal slippery to hold,
but, more important, it would probably “gum up” the
mouth or claws of a predator due to its adhesive qualities.
Predators may also find it repulsive to the taste. What-
ever the particular defense mechanism, it must be effect-
Page 461
ive for I have seen no mutilated specimens or evidence
of attack as was so often seen in other species.
Dendronotus gracilis BABA, 1949
Dendronotus gracilis Baba, 1949, Opisthobr. Sagami Bay, pp.
87 - 88, 167; fig. 109; plt. 35
Taxonomic Remarks
To describe the internal and external structure and eco-
logy of Dendronotus gracilis, 1 have used the description,
text figures, and plates given by Basa (1949) supplement-
ed by information from Dr. Michael Miller. Basa’s data
have been rearranged to suit the format I have used in this
paper.
The radula of Dendronotus gracilis is not very different
from that of D, albus or the new species, although that of
the new species usually has more denticles on the lateral
teeth. The number of cerata (4 pairs) and their branching
pattern (very few secondary branches) in D. gracilis is
markedly different from that of D. albus (5-7 pairs of
cerata with considerable secondary and tertiary branch-
ing). Also the coloration of D. gracilis is distinctly differ-
ent from that of any other species of Dendronotus.
Neither the reproductive system nor any aspects of ecolo-
gy are mentioned by Basa (1949).
Body Dimensions, Texture, and Apertures
The largest specimen of this limaciform, relatively “deli-
cate” species was 2.5cm L (Basa, 1949). Miller (pers.
comm.) reports that his largest specimen is 0.6cm L.
Width and height are not given in either case.
The anus is on the right side between cerata no. 1 and
no. 2. Below and just anterior to the first right ceras lies
the genital aperture. The cardiac prominence is not men-
tioned. According to Basa (1949), the body is smooth,
but Miller (pers. comm.) says the body is covered with
small conical tubercles.
Foot
There is no mention of the foot by Bapa (1949). The
figure (Basa, op. cit., plt. 35) shows the dorsal aspect of
the animal so no information regarding the foot can be ob-
tained from it.
Color
The body color is bluish-white and the cerata, crown and
veil papillae are opaque white. The clavus of the rhino-
Page 462
phore is dark yellow (Basa, 1949) or opaque white (Mil-
ler, pers. comm.). This color pattern is somewhat akin to
that of Dendronotus albus, but the rest of the “coloration
of [D. gracilis] is unmistakable” (Basa, op. cit., p. 167).
About 15 “yellow ocelli, each with an orange spot in
the centre, are irregularly scattered on the back [dorsum]”
(Basa, loc. cit.). The color plate also shows what may be
4 patches of white pigment in a mid-dorsal medial line ex-
tending from the posterior pair of cerata to the tip of the
tail (these may also be the 2 “accessory cerata” mentioned
below; see Cerata). Miller (pers. comm.) noted 5 longi-
tudinal rows of yellow spots: 1 mid-dorsal, 2 dorso-lateral,
and 2 lateral.
Cerata
Basa (1949) mentioned 4 pairs of short cerata while
Miller (pers. comm.) found 5 pairs on the 0.6 cm animal.
These divide into 2 or 3 main branches in what appears
to be a “rosette” pattern (Figure 4b). There are fewer
secondary branches than in any other species of Dendro-
notus and tertiary branches appear to be lacking com-
pletely. Posterior to the last pair of cerata are 2 “accessory
cerata,” the size and branching pattern of which is not
described.
Rhinophore and Clavus
There is neither a lateral papilla nor a posterior crest
(such as Dendronotus iris has) on the round rhinophore
stalk. The clavus is perfoliate with 8-17 leaves (Baza,
1949, text fig. 109b; Miller, pers. comm.). There are
4-6 simple or branched, approximately equal, but short
crown papillae around the relatively large clavus.
Head, Lips, and Veil
Basa (1949, p. 167) described 4 relatively simple, ap-
proximately equal veil papillae. However, the diagram
(text figure 109a) shows a third pair of short, simple
papillae between the medial (no. 1) and lateral (no. 3?)
veil papillae. No lip papillae are mentioned or dia-
grammed.
Jaws
The jaw is about 2.5 mm L or about a tenth the length of
the animal. The long axis of the dorsal process is at nearly
right angles to the long axis of the body of the jaw (Basa,
1949, text fig. 109d). This is a greater angle than seen in
any of the other species of Dendronotus and may be one
THE VELIGER
Vol. 12; No. 4
of the more useful taxonomic characters of this species.
The masticatory process of the jaw sports about 40
hook-shaped denticles along the masticatory border.
Radula
The radula formula is 41 (8-1-8) in the one specimen
Basa (1949) examined. On each side of the median tooth
cusp there are 17 - 20 relatively small denticles decreasing
in size towards the apex. The cusp is quite low, being
about 23 as wide as high (Basa, op. cit., fig. 109f).
Of the 8 lateral teeth, the medial 6 have cusps and
denticles while the outer 2 have neither. The cusp of the
no. | lateral tooth is about equal in size to the 4 denticles
of that tooth. In teeth nos. 2 - 4 the cusp increases in size
and then decreases in nos. 5 - 6. On teeth nos. 2 - 6, there
are 7 - 9 denticles which are largest on no. 2 and decrease
progressively to become mere serrulations on no. 6.
Reproductive System
No mention is made of the reproductive organs by BABA
(1949). A detailed description of the system would be
useful in more securely identifying the status of this
species within the genus.
Geographical Distribution
Basa’s one specimen was collected in August, 1939, from
160m at Amadaiba, Sagami Bay, Japan. Miller (pers.
comm.), dredging from 9 - 24 m in New Zealand waters,
collected 2 specimens on hydroids growing on Glycimeris
valves.
Dendronotus subramosus MacFarLanp, 1966
(Plate 64, Figure 33; Text figures 3, 4, 6, 19 - 21)
Dendronotus subramosus MacFartanp, 1966, Mem. Calif.
Acad. Sci. 6: 265 - 270; plt. 40, fig. 3; plt. 46, figs. 5 - 8;
plt. 47, figs. 3-7; plt. 49, figs. 1-3; plt. 50, fig. 2
plt. 52, figs. 1-2
Taxonomic Remarks
Except for the original description, no information con-
cerning Dendronotus subramosus has been published.
Marcus (1961, p. 34) includes, under D. frondosus,
animals which “... are translucent grey with two brown
stripes on either side between the insertions of the cer-
ata ....” but these are probably D. swbramosus. Ecolog-
Vol. 12; No. 4
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Page 463
ical as well as morphological data lead me to consider D.
subramosus a valid species, distinct from D. frondosus.
Body Dimensions, Texture, and Apertures
The limaciform, moderately “heavy” body, is laterally
compressed. The rounded dorsum merges into the vertical
sides with no distinct dorso-lateral boundary save the
cerata (Figure 3). The largest animal that I have found
was 6.5cem L & 18cm H X 0.9cm W while that of
MacFartanp (1966) was 4.0cm L & 0.8cm H X 0.55
cm W. The mode was about 2.8 - 3.0 cm L (0.3 - 3.9 cm)
while the mean length, 2.4 cm, agrees well with that of
MacFartanp’s 2.5 cm. Some others measured were: 0.9
cm L X 0.15cmH & 0.2cm W; 24cm L & 0.4cm H
<x 0.35cem W; 23cm L X< 0.5cm H & 0.35 cm W.
The anus is usually inconspicuously located about half
way along the outermost dark brown line between no. 1
and no. 2 right cerata (see Color, below). The anal
papilla, if present, may be capped by yellow pigment.
The genital openings are from 4 to 2% the way up
the right side, just anterior to the base of the first ceras.
They are relatively inconspicuous except that the tip of
the preputium often has a ring of pale yellow pigment
around it.
The cardiac prominence is usually quite large, but in
a few animals it was indistinct. To some degree, the size
can be altered in the individual.
The body and appendages may be smooth, but more
often they are covered with relatively small, closely-set,
sub-conical to conical papillae. These papillae range in
size up to 3mm, the largest ones usually situated on the
dorsum, especially the cardiac prominence, while the
smaller ones are on the sides and the appendages. Often
patches of lemon-yellow or white pigment adorn the
papillae.
Foot
The long, narrow foot is bluntly rounded anteriorly and
comes to a short, blunt point posteriorly. The sole of the
foot is a translucent pinkish-white. When crawling, the
edge of the foot is often flared laterally, especially in the
posterior half. In a specimen 2.3 cm long, the foot was
2.00cem L & 0.25cm W while, in a 4.0cm specimen,
MacFarianp (1966) found the foot to be 3.7cm L &
0.35 cm W.
The foot, the edges of which are able to fold together
very easily, is well adapted for clinging to the hydroid,
Aglaophenia struthionides, upon which Dendronotus sub-
ramosus is usually found. It is only with much difficulty
that the animal can be shaken from its perch; a most
useful attribute considering that it often occurs in areas of
considerable current action.
Color
The ground color varies from white (13 specimens)
to dark brown (3 specimens) with a few being pale yel-
low, deeper yellow or bright orange (9 specimens). Most
are reddish-brown (41 of 66) (Plate 64, Figure 33). A
liberal sprinkling of pale lemon-yellow or orange spots is
densest on the tips of the appendages, less so on the dor-
sum and sparsest on the sides. There are also variable
numbers and combinations of tiny brown, red-brown,
gold, green, and white spots intermingled with the former.
A distinct pattern of 4 light to dark brown, longi-
tudinal lines is usually seen in the intervals between the
dorsal appendages. The outermost line on each side con-
nects the lateral base of the rhinophore stalk with the
lateral base of each successive pair of cerata and carries
on to the tip of the tail. Likewise, the inner line on each
side connects the medial base of the rhinophore stalk to
the medial base of each successive ceras and again is ex-
tended towards the tail where all 4 lines merge. These
lines, particularly the posterior or medial ones, or both,
may be found wanting in lighter colored animals. The
areas between the lateral and medial dark bands are usu-
ally devoid of brown pigment but often are liberally
covered with yellow or white spots.
Cerata
There are usually 5 (3-6) pairs of stout, upright cerata
spaced at posteriorly decreasing intervals along the dorso-
lateral borders of the body with an occasional single ceras
at the posterior end (Figure 3). The size and degree of
branching of the cerata also decreases from anterior to
posterior.
The thick stalks subdivide about halfway up the total
height of the cerata in a “rosette” pattern (Figure 4b).
Determining the number of main branches may be difh-
cult, but there are usually 3 (2-4) main branches in
each ceras of the first pair, 2 (2-4) in the second pair,
2 (1-4) in the third pair, 2 (1-4) in the fourth pair,
and 1 (1-2) in the fifth pair. The postero-medial bran-
ches are usually the longest and most branched. The
lateral branch, found arising from the base of the ceras
in all the other species is missing in this animal. There
are relatively few of the stout secondary branches and
Page 464
even fewer of the very short, simple tertiary branches
making the ceras less delicately and complexly branched
than is common in the genus.
Only the first pair of cerata, as well as the rhinophores,
receives hepatic diverticula which arise from the anterior
lobes of the liver; there are no diverticula from the
posterior lobe.
Rhinophore and Clavus
The rhinophore (Figure 3), stout, erect, and inclined
slightly forward when the animal is crawling, completely
lacks a lateral papilla.
The perfoliate, bluntly conical clavus carries 9 - 14
(3 - 14) leaves (8 according to MacFartanp, 1966). The
number appears to be related to size, the small animals
(0.3 -0.9cm L) having 3-6 leaves and the larger ones
(3.7-6.5cm L) having 12 - 14 leaves. These leaves are
relatively shallow (narrow in breadth) and show little
alternation in size although they do become smaller near
the top. The coloration of the clavus matches that of the
rest of the body.
The campanulate rhinophore sheath has 5 (3 - 7) short,
blunt crown papillae. About equal in size, the papillae
are not much taller than the extended clavus; the postero-
medial is the longest by about 14. A few small bran-
ches may be found on the longer crown papillae, but the
rest are simple.
Head, Lips, and Veil
On a very distinct horseshoe-shaped veil, there are usu-
ally 2 pairs of veil papillae (Figure 3). There may occa-
sionally be a third pair (MacFar.anp, 1966) or, more
commonly, an extra papilla on one side or the other.
The medial pair (no. 1) of veil papillae is the longest
and normally points straight ahead when the animal is
crawling. About 4 to 2 as long as the medial pair, the
lateral pair (no. 2) is usually directed at right. angles
from the body. If a third pair (or one of the pair) is
present, it usually is a mere tubercle positioned laterally
to no. 2. Pairs no. 1 and no. 2 are both stout, with only
a few short, blunt branches being given off, usually from
the distal half of the papillae.
MacFarianp (1966) neither mentions nor figures any
lip papillae, but I found 2 (1-4) in about 50% of the
animals examined. They are short and simple, arising at
the posterior end of the lips.
Jaws
The jaws (Figure 19) are a striking reddish-brown,
darkest on the hinge and middle of the body, and
THE VELIGER
Vol. 12; No. 4
becoming lighter around the edges of the body and on
the dorsal process.
The main body of the mandible, thin, mytiloid, and
ovate in form, is slightly more than half as wide as long.
Cc 1.0mm D
-——————_
Figure 19
Dendronotus subramosus jaw
B. Ventral view C. Inside lateral view
D. Outside lateral view
A. Dorsal view
The dorsal process, curving backward at about 65°
from the longitudinal axis of the body, is about 4 as long
as the body, shallowly grooved dorsally, and rounded at
the tip. The triangular expansion, joining the proximal
half of the dorsal process to the body, bulges outward
only moderately.
According to MacFartanp (1966), the short, curved
masticatory margin is armed with about 120 closely set,
transverse, smooth, crescentic plates, narrowest at the
hinge and largest at the distal end. ‘The masticatory
process is joined to the body by a thin, inward-bulging
plate.
Radula
The radula formula is 59-72 (5-7:1-5-7) according
to MacFarianp; I found it to be 54 - 62 (2-5°1-2-5)
in the 5 specimens I examined (Figure 20). The medial
tooth has a small cusp, the width: length ratio being
about 2:1 (Figure 6 for measurements). Seven to 10
(2-12) long, heavy denticles, becoming smaller towards
the apex, adorn each side of the cusp. The oldest teeth
often have only 2 denticles, only slightly smaller than the
main cusp and set off from it. Most of the rest are like
those shown in Figure 20, although, in the youngest
medial teeth, the cusp may be crescent-shaped.
Vol. 12; No. 4
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Page 465
The lateral teeth are flat, often irregularly shaped
plates. At the anterior end of the radula, there is usually
only 1 tooth on either side, but this number increases to
about 3 (2-4) a third of the way along and by the half
to two-thirds point, the full complement of 5 (4-7)
teeth is usually present. The pointed, spike-like cusp is
Figure 20
Dendronotus subramosus radula
Rows 34 and 35 in a radula with 53 rows of teeth
variable in length, being short on no. 1, longest on no. 2
and no. 3, shortest on no. 4, and missing on nos. 5 to 7.
Teeth nos. 5 - 6, when present, are usually narrow, lacking
both cusps and denticles, while no. 7 is a mere rudiment.
The denticles, moderately sharp and long, are usually
variable in shape and orientation. Tooth no. 1 has 3
(2-6) denticles about equal to the cusp in size; no. 2
has 5-6 (2-8) long denticles; no. 3 has 3-4 (2-8)
shorter denticles; no. 4 has 2 (0-4) short denticles; and
nos. 5 - 7 have none.
Reproductive System
The hermaphrodite duct joins the white to light pink
hermaphrodite gland to the short, thick, crescentic am-
pulla (Figure 21). The comparatively long spermoviduct
arising from the ampulla bifurcates to form a short
proximal portion of the vas deferens and a somewhat
longer thin-walled oviduct, contrary to what MacFar-
LAND (1966) figures.
The prostate, composed of 6 - 7 alveoli (10 - 12 accord-
ing to MacFartanp) forms a single circle around the
duct. The convoluted, fairly short, distal portion of the
vas deferens tapers considerably from a wide, thick-walled
duct near the prostate to a very narrow tube where it
enters the base of the penis. The long, thin penis, exten-
sively coiled in the thin-walled preputium, is tapered
throughout its whole length.
DVD PE
ID
1.0mm Tent
Figure 21
Reproductive system of Dendronotus subramosus (exploded view)
AMP = ampulla BC = bursa copulatrix
DVD = distal vas deferens FC = (to) fertilization chamber
FGA = female genital aperture FGM = (to) female gland mass
HD = hermaphrodite duct ID = insemination duct
OV = oviduct P = preputium PE = penis
PR = prostate PVD = proximal vas deferens
SO = spermoviduct SR = seminal receptacle VA =vagina
VE = vestibule
The vagina is slender and somewhat convoluted before
it gives rise to the short, stalked, spherical seminal recep-
tacle. This in turn gives rise to a short, thick insemination
duct. The small, sac-like bursa copulatrix is located near
the opening of the vagina into the vestibule.
Geographical Distribution
In California, MacFartanp (1966) reports Dendronotus
subramosus in the intertidal from Humboldt Bay to New-
port Bay and especially in Monterey Bay. Lance (pers.
comm.) has collected the species in the Macrocystis sp.
canopy at 18m just south of South Island, Islas Los
Coronados, Mexico. Locally, it has been collected from
San Juan Channel, Edwards Reef, Lonesome Cove, Low
Island, and Peavine Pass, all sub-tidally.
Ecology
MacFartanp (1966) found Dendronotus subramosus
most commonly in tide pools during the summer, but it
Page 466
was present all year long. Locally, I found it most com-
monly from June to November, but this may reflect the
timing of collecting trips.
I collected Dendronotus subramosus in 15-30m of
water (except for 5 specimens dredged from 120 m in San
Juan Channel), but I have not seen them intertidally
despite many collecting trips. All the collection sites are
swept by fast currents, up to 4 knots or more. The bottom
is generally composed of gravel, shell, and sand, over
which are scattered many small to large stones. This type
of bottom normally supports an extensive hydroid fauna
including Aglaophenia struthionides upon which D. sub-
ramosus are found. In the laboratory, they showed a pref-
erence for Aglaophenia over any other substratum about
90% of the time. The food of this species consists of
whole hydranths of A. struthionides as well as sections of |
the coenosarc that are drawn through holes rasped or
chewed in the perisarc.
Animals with a mature hermaphrodite gland were
found from October to November, 1966 and one in Feb-
ruary, 1967. During this period of time (October to No-
vember) about 10 egg masses were deposited in the aqua-
ria. At the same, in the field, many egg masses were
seen on the top third of the Aglaophenia fronds. During
June and July, 1967, about 150 egg masses were seen at
Low Island and Edwards Reef, again on Aglaophenia.
This placement of the egg masses seems disadvantageous
in that they are probably visible to any hungry fish. On
the other hand, it exposes the egg mass to the current,
thus keeping it relatively clean and facilitating the dis-
persal of the planktonic veligers. Also, those smaller,
benthic predators that might otherwise eat the eggs
probably do not detect them when the eggs are high off
the bottom.
The white to pink egg masses are laid in a loosely coiled
string which itself is arranged in a loosely spiralled
clump and attached to the substratum by a capsule-free
mucous sheet. In the 4 cases timed, the veligers hatched
15 - 18 days after oviposition.
Dendronotus subramosus on Aglaophenia is most cryp-
tic and it is only with careful searching that one is able
to find it amongst the branches of the hydroid, particular-
ly while diving. Only the very light colored ones are
visible against the dark brown of the hydroid. The nudi-
branch is able to swim clumsily when disturbed. It has
no other obvious means of defense, so perhaps its cryptic
coloration is its main means of survival.
THE VELIGER
Vol. 12; No. 4
Dendronotus albus MacFartanp, 1966
(Plate 64, Figure 34; Text figures 2, 4 - 6, 22 - 24)
Dendronotus albus MacFaruanp, 1966, Mem. Calif. Acad.
Sci. 6: 275 - 279; plt. 40, fig. 1; plt. 46, figs. 1 - 4; plt.
47, figs. 8- 11; plt. 48, figs. 7 - 8; plt. 49, fig. 5; plt. 50,
fig. 4; pit. 51, figs. 6-7
Taxonomic Remarks
The description of this species is based on the extensive
descriptions of each of 81 animals collected between Sep-
tember 14, 1966 and January 2, 1967 at various points
around San Juan Island. About 100 more animals were
examined to see how much, if any, variation there was
from the first group.
The reader is referred to the section on Taxonomic
Remarks under the new species of Dendronotus for a
discussion of the difficulty of separating it from D. albus.
Also included is a discussion of why I have considered the
two species distinct.
Body Dimensions, Texture, and Apertures
The long, slender, moderately compressed body is quite
“delicate” as might be expected in an animal this small.
The rounded dorsum tapers posteriorly to a short, narrow,
sharp-pointed tail, and merges laterally with the nearly
vertical sides with no hint of a border save the cerata
(Figure 2).
The largest animal was 4.0 cm L, and the shortest was
0.4 cm L. MacFartanp (1966) reports a maximum size
of 3.0cmL X 0.5 cm H X 0.3 cm W. Most of the animals
were between 1.5 and 2.5cm L with an average of 1.9
cm L. Some representative measurements are: 14cm L
«< 0.4cmH & 0.25cm W;1.8cmL X 0.3cem H & 0.35
cm W;2.3cmL X 0.35cem H «0.3 cm W; 2.9cemL X
0.3cm H X 0.25cm W. The adult size is thus much
smaller than that of the new species of Dendronotus.
The anus is borne on a small papilla located halfway
between no. 1 and no. 2 pairs of cerata on the right
side. Often, the top or the sides of the papilla, or both,
are encrusted with an opaque white pigment.
The genital orifices are situated about 3 of the way up
the body just anterior to the base of the first right ceras.
The distal end of the preputium often protrudes and
bears a white pigment spot.
The cardiac prominence may be markedly elevated
and it can vary in an individual. Most of the body surface
Vol. 12; No. 4
THE VELIGER
Page 467
is smooth, but about 50% of the animals have few to
many small, bluntly conical papillae on the dorsum. They
are largest and most common on the cardiac prominence.
The yellow pigment which normally appears on these
papillae forms what looks like a small peg passing into
the dermis.
Foot
The long, narrow foot, about 8 - 10 longer than wide,
is bluntly rounded anteriorly and tapers fairly sharply in
the posterior quarter to a point. In an animal 2.65 cm L,
the foot was 2.4cm L x 0.3 cm W. When the animal is
crawling, the foot flares laterally, especially in the pos-
terior third. The sole of the foot is translucent white.
Dendronotus albus can cling so tightly to a hydroid
that it is extremely difficult to shake it loose and one must
resort to peeling the nudibranch away from the stalk of
the hydroid. This clasping ability enables the nudibranch
to remain on the hydroid, Thuzaria argentea, in the face
of some very strong currents.
Color
The translucent body is usually white although a few
pale cream or pink-brown specimens were seen (Plate 64,
Figure 34). The color of these latter is due to a dense
sprinkling of tiny, opaque, white, yellow, or pink-brown
spots over the dorsum and appendages. One animal was
a medium red-brown color, palest on the sides and darkest
on the dorsum and cerata. The internal organs show
through as a red-brown or white mass, depending on the
animal’s sexual maturity.
The pattern of encrusting, opaque, dead-white pigment
consists of solid white lines or broken patches in varying
amounts on the postero-medial (or occasionally antero-
medial) face of the cerata’s main branches; the posterior
face of the rhinophore stalk and the posterior crown pa-
pillae; and the dorsal surface of the veil papillae. A
dorso-medial white line begins between the second and
fifth pairs of cerata and extends to the tip of the tail.
Anteriorly, it may be broken up into irregular patches.
The cerata are striking in many specimens. The brown
hepatic diverticula become a very dark, rich brown
about 4 to $-way up the branch. From about 4 to 2 of the
way up, this brown merges with a very beautiful metallic
orange or copper pigment, and this in turn merges with
the opaque, dead-white pigment near the tip. In a few
specimens, the orange or copper pigment was missing,
being replaced by the white while a few others lacked
both pigments and the brown of the diverticula continued
to the top. This elegant pigmentation normally only
occurs on those branches containing hepatic diverticula,
but all the other main branches do have the white pig-
ment on the tips.
Cerata
There are 5-7 (4-8) pairs of tall, delicate, moderately
branched cerata arranged at posteriorly decreasing inter-
vals on either side of the dorsum (Figure 2). The height
of the cerata also decreases posteriorly and the last pair
is often less than a quarter as tall as the first pair. In
almost half of the animals, there are one or two short,
simple, unpaired posterior papillae.
The main branches are long and slender with a few
relatively long secondary branches. In contrast to the new
species of Dendronotus there are many small, almost trans-
parent tertiary patches giving the cerata a “bushy” ap-
pearance.
The branching pattern is “fan-shaped” (Figure 4a)
with 3 main branches in the first and second pair of
cerata, 2-3 in the third pair, 2 (1-3) in the fourth
pair, 1-2 in the fifth pair, and one in pairs 6 to 8. In
those pairs with 3 branches, the lateral branch, from 4 to
2 as long as the medial, arises from the base of the ceras.
The main stalk then branches a short distance above this
into 2 almost equal branches. When there are only 2
main branches, the medial is from 2 - 4 longer than the
lateral which again arises from the base of the ceras. This
lateral branch decreases in size from anterior to posterior
and, in the last two pairs of cerata, it is usually missing.
The hepatic diverticula extend into the 2 medial
branches (trifid cerata) or the single medial branch (bi-
fid cerata) in the first to fourth (1-6) pairs of cerata as
well as the rhinophore stalk and posterior crown papillae.
The anterior lobes of the liver give rise to the diverticula
of the first pair of cerata and the rhinophore while the
posterior lobe gives rise to the rest of the diverticula.
Rhinophore and Clavus
The erect, delicate rhinophore bears few branched proc-
esses, much like the new species of Dendronotus.
The lateral papilla, varying in length from a mere bump
to a relatively long, slender papilla, is usually unbranched
or, at most, bears a few small, simple papillae. It arises
about 4 to halfway up the rhinophore stalk.
The conical, perfoliate clavus bears 13-17 (12-22)
deeply-cut, alternating leaves (12 - 14 according to Mac-
FarLAND, 1966). These are usually peppered with pinkish
spots that impart a distinctly tan color to the clavus.
Page 468
Often, the distal half of the clavus is bent forward 90
to 120° from the vertical.
The slender, unbranched crown papillae are directed
obliquely upward, but are so tall that they tend to droop
outward. The anterior 3 are shortest and about equal in
length. The postero-medial one is about 3 (2-4) as
long as the anterior ones and often stands taller than the
rhinophore stalk. The postero-lateral papilla is often
nearly as long as the postero-medial and usually at least
2 longer than the anterior papillae.
Head, Lips, and Veil
These structures are remarkably similar in size and loca-
tion to those in the new species of Dendronotus. The 3
pairs of long, slender veil papillae are usually simple
(Figure 2). Sometimes they have a few short, simple
secondary papillae and, only rarely, very few tertiary
branches. ‘The medial pair (no. 1), directed forward and
slightly upward, is the longest. The middle pair (no. 2)
is about 4 to 4 as long as the medial pair, originates vent-
ro-lateral to the base of no. 1 and is directed ventro-
medially, in a hook-like fashion, toward the mouth. About
= Of the length of no. 1, the lateral pair is often split
into 2 rami with the lateral ramus about + to 2 as long
as the medial ramus. Both rami are directed upward and
laterally at 70 - 90° from the long axis of the body.
There are 2-4 (0-8) long, thin, blunt, unbranched
lip papillae located on the posterior border of the plicated
lips and lateral to the mouth. Being very sensitive, these
papillae retract rapidly if touched, thus possibly account-
ing for the wide range in numbers observed.
Jaws
The jaw (Figure 22), very small and fragile, is neverthe-
less relatively large for an animal of this size. In a pre-
served specimen 1.2cm L (1.5cm L when alive), the
jaw was 0.22cm L & 0.19cm W & 0.15 cm H (Figure
5 for measurements) about a seventh as long as the ani-
mal. The body of the jaw is a horn yellow while the hinge
area, masticatory process, and most of the dorsal process
are a darker brown, gradually fading to horn yellow near
the body. The right and left halves of the jaw diverge
posteriorly more than in any other species of the genus;
this is even more pronounced when all the musculature
is removed.
The shallowly convex, nearly oblong body is about 2.2
times as long as wide, and weakly convex at the posterior
border. The moderately curved, strong dorsal process,
inclined at 50-60° from the long axis of the body, is
about 0.43 x as long as the body. A strongly convex lateral
THE VELIGER
Vol. 12; No. 4
Figure 22
Dendronotus albus jaw
B. Ventral view C. Inside lateral view
D. Outside lateral view
A. Dorsal view
expansion joins the proximal half of the dorsal process
to the body.
The short masticatory process, inclined at about 80°
from the long axis of the body and strongly curved dis-
tally, is joined to the body by a strong concave expansion.
The features that distinguish this jaw from that of
the new species of Dendronotus are the shape of the body
and the degree of divergence posteriorly of the 2 halves.
Radula
In the description of this species, a series of 15 radulae
was examined and another 25 radulae were examined to
check for additional variation. The resultant radula for-
mula is 32-38 (6-8-1-6-8) (Figure 23). In all the
specimens, the radula teeth were colorless. MacFARLAND
(1966) reports a radula formula of 36 - 38 (7 -9- 1-7-9)
and infers that all the median teeth are light yellow.
The bluntly pointed, denticulated cusp of the median
tooth is relatively smaller than in the new species of
Dendronotus; the width : height ratio varies from 1.8 - 2.0
(1.7-2.4) in the oldest teeth to 1.6-1.9 (1.1-1.9) in
the newer teeth (Figure 6). Along each side of the
median tooth cusp are 11 - 14 (7-17) denticles with the
average being 13. These are small, relatively sharp, more
or less regularly spaced, and usually increasing slightly in
size towards the apex. In a very few animals, the dent-
icles are missing or the edge of the cusp is very jagged
from pieces being broken out.
Vol. 12; No. 4
The tall lateral teeth are relatively stout, and the inner
ones bear long, curved cusps inclined about 20° toward
the midline. In some of the innermost lateral teeth, the
cusp is just slightly larger than the denticles, but more
often it is about 2 longer. In the next 3-4 teeth, the
cusp becomes progressively longer while teeth no. 5 (and
no. 6) have a shorter spike-like cusp. In nos. 6 to 8, the
cusp is usually missing.
Figure 23
Dendronotus albus radula
Rows 25 and 26 in a radula with 40 rows of teeth
The proximal, lateral margin of the lateral tooth cusp
bears from 3 - 8 relatively tall, stout, sharp-pointed den-
ticles at nearly regular intervals. The 3 innermost laterals
bear 4-6 (3-8) denticles while the next 2 bear 4-7
(3-9) denticles and no. 6, when it has denticles, carries
4-7 of them. The size and number of these denticles are
rather distinct from those in the new species of Dendro-
notus.
Reproductive System
The thin hermaphrodite duct empties into a very wide,
short, crescentic ampulla which suddenly narrows to a
long spermoviduct (Figure 24). This divides to form a
short, wrinkled oviduct and a very short proximal portion
of the vas deferens. The prostate, comprised of 12-15
large, irregularly shaped alveoli arranged around the vas
deferens as a concentric disc, is much smaller than that
of the new species of Dendronotus. From it, the relatively
short, quite narrow, weakly convoluted distal portion of
the vas deferens arises. It tapers slightly immediately prior
THE VELIGER
Page 469
to entering the penis. The moderately long, narrow penis
is tapered to a point.
AMP “Hp OR
Figure 24
Reproductive system of Dendronotus albus (exploded view)
AMP = ampulla BC = bursa copulatrix
DVD = distal vas deferens ED = ejaculatory duct
FC = (to) fertilization chamber FGA = female genital aperture
FGM = (to) female gland mass HD = hermaphrodite duct
ID = insemination duct OV = oviduct P = preputium
PR = prostate PVD = proximal vas deferens
SO = spermoviduct SR = seminal receptacle
VE = vestibule
PE = penis
VA = vagina
The distal portion of the vagina passes through a very
muscular mass, with unknown function, before it enters
the vestibule. The long, flaccid, sac-like bursa copulatrix
is located at the distal end of the vagina right over this
mass. The rest of the vagina is moderately long and quite
narrow, emptying finally into a stalked, spherical seminal
receptacle. The insemination duct, arising from the base
of the seminal receptacle, is very short, wrinkled, and
transparent.
The major differences between Dendronotus albus and
the new species are in the size and shape of the prostate,
ampulla, vas deferens, and seminal receptacle.
Geographical Distribution
Dendronotus albus has been collected at a number of
points around San Juan Island: Brown Island (10),
Collins Cove (8), Lonesome Cove (3), and Edwards
Reef (200). The only other area was Albert Head, Vic-
tonasp bara (2))e
MacFarianp (1966) collected Dendronotus albus at
Monterey Bay. Lance (pers. comm.) reports collecting
one specimen each at 3 places: the lower intertidal of
Moss Beach (June 13, 1964) ; at 25 m in the Macrocystis
sp. canopy near Scripps Canyon (July 24, 1967) ; and at
18m in the Macrocystis sp. canopy south of the South
Island, Islas Los Coronados, Mexico (June 22, 1961).
Page 470
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Vol. 12; No. 4
Ecology
From September, 1966 to January 1967 and during
March, 1968, the animals were seen at depths of 20 - 30 m
on rocky bottoms swept by strong currents most of the
time. At Edwards Reef, the substratum consists of stones
up to 18 inches across thickly scattered over a shell-gravel-
sand base. These areas support dense, extensive hydroid
populations of which Thuzaria argentea is often the most
common species. Usually, the nudibranch was found
clinging to the tip of the hydroid, and even careful
searching failed to turn up more than a few animals
crawling on the rocks. At Collins Cove where T: argentea
is rare, the nudibranch was found on Hydrallmania dis-
tans. Laboratory observations suggest that Dendronotus
albus, given a choice, will nearly always (95%) choose
T. argentea over any other substratum.
Although no data are available on the mechanics of
feeding, Dendronotus albus does eat Thuzaria argentea,
normally confining itself to the distal portions of the
stalk. Apparently it does not ingest the perisarc as this is
not found in the fecal pellets.
The white egg masses are usually deposited at the
tip of the branch of Thuiaria argentea, thus raising the
eggs off the bottom and facilitating dispersal of the veli-
gers as well as keeping the eggs clean. Many egg masses
were deposited in the field and in the laboratory during
October and November, 1966. No record of the number of
eggs per capsule or the hatching time was kept. Copulation
and well developed gonads were seen during March, 1968.
The apparent reproductive isolation of the new species of
Dendronotus and D. albus is mentioned below.
Both the nudibranch and its eggs are very conspicuous
on the tips of the hydroids, making them easy to collect
and probably easily seen by fishes. They have no obvious
defense against predators, particularly fish, but the large
number of mutilated specimens suggests that many are
attacked and subsequently escape. Perhaps they have a
distasteful epithelial secretion as do many other nudi-
branchs (Epmunps, 1966; THompson, 1960b).
Dendronotus albus is able to swim quite well but its
smali size limits the absolute distance covered. Possibly,
with the aid of currents, swimming may be an effective
means of transport for the adults.
Dendronotus diversicolor RoBILuiarD, spec. nov.
(Plate 64, Figures 35, 36; Text figures 4 - 6, 25 - 28)
Taxonomic Remarks
Marcus (1961, p. 34) includes under Dendronotus fron-
dosus specimens that “ are white with orange-
yellow points on the principal branches of the append-
ages,” but these are probably D. diversicolor.
On the basis of MacFarLanp’s (1966) description of
Dendronotus albus it is difficult to separate D. albus from
D. diversicolor. However, after examining many speci-
mens, I was able to detect some obvious and some subtle
internal and external characters that are consistently
different in the two species. There is also some ecological
evidence that attests to their uniqueness. These are indica-
ted with a fuller description under the appropriate head-
ings in both D. albus and D. diversicolor.
The most consistent morphological differences are in:
the body size at maturity; the number of pairs of cerata;
the body texture; the coloration and its pattern; the
number and location of the hepatic diverticula; the denti-
culation of the radula; the overall shape of the jaws; and
the proportions and shapes of the organs of the reproduc-
tion system. A thorough knowledge of the diagrams and
descriptions of the above characters should allow the
reader to distinguish one species from the other.
It could be argued, though with little validity, that
the variation of the above listed characters is within the
range of variation for a hermaphroditic gastropod. How-
ever, there are other factors which are not so easily
reconciled.
The “ceratal cores” or hepatic diverticula extend into
3-5 pairs of cerata in Dendronotus albus and only into
the anterior 2 pairs in D. diversicolor. It seems unlikely
that those in the posterior 1-3 pairs should suddenly
regress when the animal grows longer than about 3 cm.
In fact, the larger D. albus (2.5-4.0cm L) have diver-
ticula in the 5" and 6" pairs of cerata as well. In the
same vein, it seems unreasonable to postulate a sudden
regression of the last 2 - 3 pairs of cerata as well as many
of the smaller branches in the anterior pairs when the
animal attains a length of about 3 cm.
Probably the most critical difference is the lack of
sexual activity between the two species. No interspecific
copulation was seen in the field or in the laboratory
despite many observations of intraspecific copulation in
the same animals. On many occasions, both species were
placed together in an attempt to induce interspecific
copulation, but with no success. Finally, Dendronotus di-
versicolor does not seem to develop mature gonads until
it is more than about 4.0 cm L, whereas D. albus may be
sexually mature and depositing eggs before it is 1.2 cm
L. It might be argued that the larger animals are 2
years old and, having reproduced once, lived through the
winter to breed again. Or, they may not have bred the
first year; instead, they may have channeled the excess
energy into growth and reproduced the second year. Mit-
LER (1962) suggests this may be true for D. frondosus.
Vol. 12; No. 4
Tuompson (1964) reports that most hydroid-eaters pass
through a number of generations in a year or at least live
only one year. Preliminary observations in the field tend
to support THompson, at least for the small species of
Dendronotus, including D. albus and D. diversicolor. It
is for these reasons that I feel justified in establishing D.
diversicolor as a new species.
The name has been chosen to draw attention to the
variable pattern of pigments on the body and appendages,
and the two color phases. The type locality is designated
as Cantilever Pier, San Juan Island, Washington (lat.
48°32’48” N; long. 123°00’18” W). The collection of
specimens from which the description is taken is deposited
in the California Academy of Sciences, Invertebrate Type
Series, Holotype no. 416. Paratypes are deposited at the
Smithsonian Institution, and the Canadian National Mu-
seum (Ottawa). ‘
Body Dimensions, Texture, and Apertures
The slender, limaciform body is relatively “delicate” for
an animal of its size, and more laterally compressed than
in the other species of Dendronotus. Because of the lack
of cerata on the posterior third of the body, there appears
to be a long tail tapering to a point. The high, rounded
dorsum merges into the vertical sides with only the cerata
to mark the junction between the two. In the present
study the largest of 25 specimens was 7.3 cm L and the
shortest was 1.5cm L while most were 3.0-6.0cm L.
Representative sizes were: 5.2cm L & 0.8cmH X 0.4
emis 5) lemme <1 OY/omi Et <7 0)5 em) Wie 19yem)
< 0.3cm H & 0.2 cm W.
The small anal papilla, situated halfway between the
first and second right cerata, is often covered with an
opaque, dead-white pigment.
The genital apertures, which may not be conspicuous,
are located about halfway up the right side just anterior
to the base of the first right ceras.
The cardiac prominence may be only slightly raised,
but more often, it is markedly protuberant. The surface
of the body is very smooth; none of the specimens showed
any sign of the papillation seen in other species.
Foot
The foot, about 10 as long as wide, is bluntly rounded
in front and tapers in the posterior third to the sharply
pointed tail. The sole is white in white animals, but is pale
pink in lilac-colored animals.
The labile foot is able to cling most tenaciously to
hydroid stems, making it nearly impossible to shake the
animal loose. To be studied, it must be literally peeled
off the stem. This degree of adhesion is extremely effective
THE VELIGER
Page 471
in allowing the animal to maintain its position despite
the swift currents that normally sweep past it. The edges
of the foot fold closely together when the animal swims.
Color
The translucent body may be either white or lilac (Plate
64, Figures 35 and 36). Fifteen animals were white while
10 were varying shades of lilac from cameo pink to pale
violet. An opaque dead-white or a striking, opaque orange
pigment, or both, may be found on the distal third of the
main branches of the cerata, the posterior 1 or 2 crown
papillae, the back of the rhinophore sheath, and the tips
of the veil papillae.
In 15 white specimens, 13 had mostly white pigment
(5 of these had a bit of orange at the very tip) and 2 had
mostly orange pigment on the cerata. Each group also had
a narrow orange or white band, respectively, proximal to
the main pigmentation. Of the 10 lilac forms, 3 showed a
predominance of white with only tiny caps of orange
and the other 7 were basically orange with a narrow band
of white proximal to the orange.
In all the specimens, there is an opaque dead-white,
dorso-medial line extending from between the last pair
of cerata to the tip of the tail.
Cerata
The 4, sometimes 5, pairs of tall, slender, sparsely branched
cerata are arranged in a single row along either side of
the dorsum with the interval between pairs decreasing
very little. The height of the first 3 pairs of cerata is
about equal while the 4" pair is about 4 to 3 as high.
If present, the fifth pair is reduced to simple, short
papillae.
The main branches are tall and slender, giving rise to
few, relatively long, thin secondary branches, usually from
the distal half of the stalk. The very few tertiary branches
are short and almost transparent.
The branching pattern is “fan-shaped” (Figure 4a)
with 3 branches in the first and second pairs of cerata,
2 (1-3) in pair no. 3, 2 (1-2) in pair no. 4, and 1 in
pair no. 5. Where there are 3 branches, the nearly hori-
zontal lateral stalk arises from the base of the ceras and,
a short distance above that, the stalk splits to form 2
erect branches. In bifid cerata, the stalk divides at the
base, giving rise to a lateral branch that is about } to 3
as tall as the medial one .
Extending into the medial 1, or more commonly 2,
main branches of the first 2 pairs of cerata as well as
into the rhinophore stalk are the hepatic diverticula. The
first 2 pairs of diverticula arise from the anterior lobes of
the liver and the last pair from the posterior lobe.
Page 472
Rhinophore and Clavus
The tall, simple rhinophore bears a slender, unbranched
lateral papilla originating between a third and halfway
up the stalk and directed at about 60° from the horizontal.
Usually, the lateral papilla is about + the length of the
stalk; in a few cases, it is reduced to a mere tubercle and,
rarely, it is missing.
The conical, perfoliate clavus, in varying shades of
white, cream, or pink, bears 16-21 deeply-cut leaves
alternating in width. The narrow leaves are usually com-
pletely hidden by the wider ones.
Directed obliquely upward from the margin of the
dilated, campanulate rhinophore sheath are 5 tall, thin,
simply branched crown papillae. The postero-medial is
about 2 to 3 longer. The shorter 3 are 4 as tall as the
rhinophore stalk.
Head, Lips, and Veil
The frontal margin of the indistinct veil bears 3 pairs of
slender veil papillae that have a few short, slender bran-
ches arising from the distal half of the stalk and running
nearly parallel to it. The medial pair, normally pointing
straight ahead, is the longest. Pair no. 2, finger-like pro-
jections about a third to a sixth as long as no. 1, originate
just ventro-lateral to the base of no. 1 and are curved
ventro-medially. The lateral pair, at right angles to the
long axis of the body, is about $ as long as no. 1.
There are 2 - 4 (2-8) simple, relatively long lip papil-
lae located on the postero-lateral margin of the outer
lips. The variation in numbers of papillae suggests that
they are able to retract when disturbed; this has been
noticed in a number of species.
Jaws
The thin, delicate jaws (Figure 25) are a transparent
pale yellow, becoming more yellow at the hinge and
masticatory process. In 1 preserved and 1 live speci-
mens, I found buccal masses measuring 0.36 cm L x 0.19
cm W X 0.20cm H (2.3cm L animal); 04cm L X
0.30 cm H (5.5 cm L; live).
The body is a convex, ovate mytiloid shape, widest in
the posterior third, and about 1.8 longer than wide
(Figure 5). The wide, curved, deeply grooved dorsal
process, inclined at about 60° to the long axis of the body,
is approximately 0.4 as long as the body. The lateral
margin of the process flares outward and toward the hinge
to form a strongly convex expansion between the dorsal
process and the body. ‘The short masticatory process is
slightly (10°) inclined posteriorly.
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Vol. 12; No. 4
C Lomm, ~~ p
Figure 25
Dendronotus diversicolor jaw
B. Ventral view C. Inside lateral view
D. Outside lateral view
A. Dorsal view
Radula
In this study, a sample of 11 animals had radula formu-
lae of 33-38 (6-9-1-6-9). I found that only the
anterior median teeth in 5 out of 11 animals were light
yellow; the rest were colorless as in other species.
The strong, pointed cusp of the median tooth is 1.4 - 1.6
(1.3-1.8) & as wide as high (Figures 6, 26). Along
either side are 13-17 (7-25) small, sometimes irregu-
larly spaced denticles usually increasing in size towards
the apex. In a few specimens, the edges were incon-
Figure 26
Dendronotus diversicolor radula
Rows 21 and 22 in a radula with 40 rows of teeth
Vol. 12; No. 4
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Page 473
spicuously serrulated, particularly on the anterior teeth,
and in others these same teeth are quite jagged where
pieces have been broken off.
The long, relatively wide lateral teeth bear elongate,
slightly curved cusps inclined medially 10 - 20° from the
long axis of the tooth. In the medial lateral tooth (no. 1)
the cusp is about equal in size to the denticles, and in no.
2 it is about 2 longer than the denticles. Teeth no. 3
and no. 4 have the longest cusps, followed by no. 5
which has a long, thin, sharp cusp. The outermost 1 - 3
teeth lack a cusp entirely and no. 9 is reduced to a thin,
narrow rudiment.
Borne on the lateral edge of the cusp are numerous
relatively long, sharp denticles. On the innermost lateral
tooth (no. 1) are 4-7 (4-12) denticles, 5-10 (4 - 13)
on no. 2, 6-9 (4-14) onno. 3 and 5-11 (2-14) on no.
4. Only rarely does no. 5 have any denticles and then only
a few.
Reproductive System
A dorsal view of the reproductive system in situ (Figure
27) shows the relationship of the genital apertures as well
Figure 27
Reproductive system of Dendronotus diversicolor
In situ right dorso-lateral view to show the position of the genital
apertures
ANT = anterior
AMP = ampulla BC = bursa copulatrix
DVD = distal vas deferens E = esophagus
FGA = female genital aperture FGM = female gland mass
HD = hermaphrodite duct I =intestine
P = preputium
ST = stomach
VA = vagina
MGA = male genital aperture
POST = posterior PR = prostate
SR = seminal receptacle
as giving some idea of the relative size that the “female
gland mass” attains in a mature specimen.
The exploded system (Figure 28) indicates size and
relationship of the components. The hermaphrodite duct
suddenly widens to form a wide ampulla which is folded
1.0mm
_-———S>
Figure 28
Reproductive system of Dendronotus diversicolor (exploded view)
AMP = ampulla BC = bursa copulatrix
DVD = distal vas deferens ED = ejaculatory duct
FC = (to) fertilization chamber FGA = female genital aperture
FGM = (to) female gland mass HD = hermaphrodite duct
ID = insemination duct OV = oviduct P = preputium
PE = penis PR = prostate PVD = proximal vas deferens
SO = spermoviduct SR = seminal receptacle VA = vagina
VE = vestibule
against itself for most of its length. The short, narrow
spermoviduct bifurcates into a short, moderately wide
proximal portion of the vas deferens, and wide, trans-
lucent, thin walled rugose oviduct.
The prostate, a flattened sphere, concentric with the vas
deferens is comprised of many (30 or more) fairly large,
irregularly shaped alveoli. Arising from the far side of
the prostate is the very short, wide, weakly convoluted
distal portion of the vas deferens. Just before entering
the penis, it tapers to a narrow tube. The penis is a short,
wide, nearly straight organ which tapers gradually to a
blunt tip.
The vagina opens into the vestibule almost at the
external orifice. The small spherical bursa copulatrix,
supported on a long stalk, is located immediately proxi-
mally to the vaginal opening. The moderately long and
narrow vagina terminates in a large, stalked, seminal
receptacle resembling a squashed ovoid in shape. The short
Page 474
but wide, translucent, wrinkled insemination duct is con-
fluent with the oviduct, the two emptying into the fertili-
zation chamber.
Geographical Distribution
From July to December, 1966, I collected Dendronotus
diversicolor at Point Caution (5), Brown Island (5),
Collins Cove (8), Lonesome Cove (9), Eagle Point (8),
all on San Juan Island, as well as at Peavine Pass (1)
and Black Rock (1), both near Blakely Island in the San
Juan Archipelago. Five animals were collected off the
Victoria Breakwater in January, 1968.
Ecology
Adults were collected locally from July, 1966 to Decem-
ber, 1966, but none were seen from December, 1966 to
March, 1967. They were collected again from May to
August, 1967,and in March, 1968. All the animals were at
15 - 20 m on rock bottoms, thickly covered with numerous
hydroid species and swept by strong currents. In the field,
they were found on Abvetinaria greenei (8), Abtetinaria
spp. (6), Sertularella tricuspidata (4), Hydrallmania dis-
tans (3), and unidentified hydroids (13). In the labora-
tory they prefer Hydrallmania distans, Abietinaria greenet
and A. amphora.
The egg mass is much like that of Dendronotus subra-
mosus in overall shape. In October, 1966, one white egg
mass was seen in the field, while in the laboratory a
lilac-colored animal deposited a lilac egg mass and 2 white
animals laid white ones. Again in May, 1967, about 10
white egg masses were seen at Lonesome Cove and Eagle
Point and 1 was laid in the laboratory. There was only
1 egg per capsule; the eggs took 8 - 13 days to hatch.
Against the dark background of rocks and hydroids,
this nudibranch, perched on the distal third of the hydroid
stem, is readily visible to a diver up to 20 feet away. The
paucity of mutilated animals suggests that they have some
effective, but as yet unknown, defense mechanism or
that they are completely devoured, thus leaving no
evidence of attack.
Like many Dendronotus species, D. diversicolor swims
successfully when it is disturbed. So far, only mechanical
stimuli have been effective in eliciting this response. As
is true for many normally benthic invertebrates that
THE VELIGER
Vol. 12; No. 4
occasionally swim, the swimming of D. diversicolor ap-
pears to be randomly rather than purposefully directed,
with the currents determining the direction of the move-
ment. Swimming may provide a means of escaping some
predators, but it may also be important in the dispersal
of the adults.
DISCUSSION
There are a number of reasons for the confusion over the
validity of the species names in the genus Dendronotus,
but the main one is the early, inadequate accounts which
lack accurate and precise descriptions or diagrams, or both
of the taxonomically important internal and external char-
acters. For example, the descriptions by GmMetin (1791)
of Doris cervina and by BercH (1879) of Dendronotus
dalli and D. purpureus leave no doubt that the animals
are dendronotids, but the species designation is difficult.
The reports of O’DonocHuE (1921) were incomplete
enough to lead OpHNeER (1936, p.1108) to say that “D.
rufus ... cannot be separated (according to the descrip-
tion) [from D. frondosus].”
Because the important external characters like the
rhinophores, cerata, lip and veil papillae, and color are
very badly distorted after preservation, many of the
earlier workers, especially BERGH, contented themselves
with describing the internal anatomy. Unfortunately,
they ignored the reproductive system, which has species-
specific characters, and concentrated on the ganglia and
nerves, digestive system, and circulatory system. All of
these systems are remarkably similar throughout the genus.
Most workers did include the radula, albeit often only
briefly, thus giving their descriptions some limited use-
fulness.
Because most of the previous descriptions of Dendro-
notus have been based on one or a few specimens, there
is no indication of the range of variation in the number,
size, shape, et cetera, of the distinguishing characters.
This “typological species concept” (Mayr, 1965) was no
small hindrance when subsequent workers tried to decide
if animals they had were forms of previously described
species or a new species. 3
OpHNER (1934, pp. 229 - 230) outlined the criteria
that he felt would make a species description more accu-
rate and useful. To ODHNER’s outline I would add only
Explanation of Plate 64
Figure 33: Dendronotus subramosus MacFarLanp, the most com-
mon color form found in the San Juan Islands x 1
Figure 34: Dendronotus albus MacFartanp. The number of pairs
of cerata_ (7) and the dorso-medial white line on the tail are
sufficient to distinguish it from D. diversicolor x2
Figure 35: Dentronotus diversicolor RoBILLIARD, spec. nov., white
form. There are only 4 pairs of cerata (CE) x 14
Figure 36: Dendronotus diversicolor RoxpiLuiarD, spec. nov., lilac
form. The dorso-medial white line on the tail of this and the speci-
men in Figure 35 are not as distinct as usual x 4
Tue VELIcER, Vol. 12, No. 4 [Ropitiiarp] Plate 64
Figure 34
Figure 33
Figure 35 Figure 36
Vol. 12; No. 4
two essential criteria. The systematist should use a large
series of animals to give some appreciation of the species’
morphological variability and consistency. He should
also have an understanding of the animal’s ecology, par-
ticularly the feeding and reproductive behavior. The
validity of a species of Dendronotus (or any nudibranch)
established on the basis of the above criteria will probably
stand the test of time.
A large sample of specimens provided the basis for
separating Dendronotus albus from D. diversicolor, and
D. rufus and D. dalli from D. frondosus. Dendronotus
albus might have been considered a juvenile D. diversi-
color, had not a long series of specimens pointed out the
consistent differences as well as the limits of morpho-
logical variation in the two species. ODHNER (1934, p.
19) states that there “... seems to be no reason for
basing specific or varietal distinctions on the shape of the
radula of this genus [Dendronotus].” However, when
many more radulae are studied, limits of the variation in
the teeth of any one species are recognized and species
distinctions can be made.
Taxonomically, a knowledge of the reproductive biolo-
gy of Dendronotus is the most important aspect of their
ecology. This has been directly demonstrated in separating
Dendronotus albus and D. diversicolor. The uniqueness
of the other species has been determined partly on the
basis of their reproductive isolation from each other.
There are definite peaks in reproductive activity, as
indicated for each species, but egg masses and sexually
mature animals can be found in small numbers at almost
any time of the year. This plus the variation in size of
animals collected at any one time suggests that reproduc-
tion occurs throughout the year.
Species of Dendronotus, like many nudibranchs, seem
to be food specialists (MILLER, 1961; THompson, 1964),
a feature that is useful in making field identifications.
There are many other aspects of their ecology, discussed
previously, which contribute to an understanding of the
species and add more dimensions to the “multi-dimen-
sional species” (Mayr, 1965).
As a last point, it seems curious that 7 of 9 species of
Dendronotus should occur along the coast of the Northeast
Pacific Ocean from Alaska to Baja California, that only 2
(possibly 3) species are found in the Atlantic Ocean, and
that only 3 species are found in the Northwest Pacific
Ocean. This may be an actual biological phenomenon
with the Northeast Pacific being the center of evolution
and radiation of the genus but it is more likely a reflection
of the collection pressure. I would expect that more spe-
cies will be found in the temperate waters of South
America and Africa where relatively little collecting has
so far been done.
OpHNER (1936) lumped all the species described from
the Atlantic under Dendronotus frondosus with the ex-
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Page 475
ception of D. robustus. However, D. elegans VERRILL,
1880 is probably a specimen of D. dalli. There may be
more species which have been incorrectly synonymized
and which further work may reveal.
Further studies of the ecology as well as geographical
distribution of the genus Dendronotus are needed to elu-
cidate the problems concerning the evolution of the genus.
SUMMARY
Nine species of Dendronotus are described or redescribed:
Dendronotus frondosus (Ascantus), D. iris Cooper, D.
robustus VERRILL, D. dalli Bercu, D. rufus O’ DONOGHUE,
D. gracilis Basa, D. albus MacFartanp, D. subramosus
MacFarianp and D. diversicolor, spec. nov.
Dendronotus venustus MacFartLanp is considered a
synonym of D. frondosus. Dendronotus dalli and D. rufus
are taken out of synonymy with D. frondosus and estab-
lished as valid species. Dendronotus elegans VERRILL is
considered a synonym of D. dalli rather than of D. frond-
osus.
The geographical distribution of the species of Dendro-
notus seems to be mainly limited to the north temperate
and arctic waters. All species but D. robustus and D.
gracilis are found in the Northeast Pacific Ocean. Dend-
ronotus robustus, D. frondosus and possibly D. dalli are
found in the Atlantic waters. Only D. gracilis has been
found in south temperate waters (New Zealand).
Extensive descriptions of each of a large number of
individuals are the basis of the species descriptions. Ex-
ternal characters used, different interspecifically and con-
sistent intraspecifically, were: the number, size, and
branching pattern of the cerata, veil papillae, crown
papillae, and lateral papillae; the number of lip papillae;
the number of leaves in the clavus; the position of the
anus and genital apertures; the size and shape of the
body; the texture of the epidermis; the color and its
patterns. Taxonomically important internal structures,
described and figured, are the organs of the reproductive
system; the radula; and the jaws. Each internal and ex-
ternal structure is defined and its function, if known, is
discussed.
Of the 7 local species of Dendronotus, 5 appear to feed
almost exclusively on various species of thecate hydroids.
Of the other 2, D. iris feeds on the burrowing anthozoan,
Cerianthus sp., while preliminary observations suggest
that D. rufus feeds part of the time on scyphistomae. The
food items of D. robustus are unknown.
Dendronotus iris is found on mud bottoms where there
is little current action. Dendronotus rufus and D. dalli are
usually in rocky areas with moderate currents while D.
albus, D. subramosus, D. frondosus, and D. diversicolor
inhabit rocky bottoms swept by swift currents.
Page 476
THE VELIGER
Vol. 12; No. 4
The local species of Dendronotus usually deposit white
to cream colored Type B egg masses (Hurst, 1967), with
1 egg per capsule in D. frondosus and D. albus, to 50 or
more eggs per capsule in D. iris. The time from oviposition
to release of the veligers ranges from 10 to 20 days.
There appears to be a limited number of defense mech-
anisms in the genus Dendronotus. Dendronotus rufus
secretes a thick, very sticky mucus which may be offensive
to predators. All species are capable of swimming and
this, combined with the currents, may carry the animals
away from danger. Varying degrees of cryptic coloration
are found in D. rufus, D. iris, D. subramosus and D.
frondosus.
ACKNOWLEDGMENTS
My sincere thanks go to Dr. Alan J. Kohn who provided
considerable encouragement and advice during the re-
search for and preparation of this manuscript. I thank
Drs. Paul Illg and Robert T: Paine for their constructive
criticism. I owe thanks to Dr. Eugene Kozloff for in-
struction in a number of photographic techniques, Mrs.
Alan Kohn for assistance with drawings, Charles Birke-
land for first bringing Dendronotus rufus to my attention.
and Mrs. Charles Lambert for translating the German
papers. I would like to extend my gratitude to Dr. Robert
Fernaid, Director of the Friday Harbor Laboratories, for
the use of the laboratory facilities where this work was
done.
For valuable information on distribution and ecology of
Dendronotus, I am grateful to Mr. Kerry Clark, Uni-
versity of Connecticut ; James Lance, San Diego; Dr. D. B.
Quayle, Nanaimo Biological Station; and Dr. Michael
Miller, University of Auckland, New Zealand. Mr. Don-
ald Wobber, Hillsborough, California, kindly provided
information on the feeding behavior of D. iris, and spon-
sored a dive to allow me to make in situ observations at
Monterey Breakwater.
Mr. Allyn G. Smith, California Academy of Sciences,
kindly provided me with specimens of Dendronotus sub-
ramosus, D. albus, and D. venustus from MacFarland’s
collections and with valuable advice and information.
Information concerning the loss of O’Donoghue’s collec-
tions of D. dalli and D. rufus was kindly provided by Drs.
D.B. Quayle, Nanaimo Biological Station; Arthur Clarke,
Jr., Canadian National Museum; and Norman Tebble,
British Museum (Natural History). The absence of
Bergh’s specimens of D. dalli in the Smithsonian Institu-
tion was confirmed by Dr. Joseph Rosewater.
I am especially grateful to my wife, Carol, for typing
the manuscript.
This work was supported by a special Scholarship from
the National Research Council of Canada and a National
Science Foundation grant to the Friday Harbor Labora-
tories.
APPENDIX I
Longitude and Latitude of Collection Sites
Locality Longitude Latitude Species of Dendronotus present
Albert Head, Victoria 123°28/36” 48°23/12” D. dalli, D. albus
Alki Point, Seattle 122°24'36” 47°34'10” D. rufus
Bellingham Bay, Bellingham 1220334 48°45’ D. iris
Brown Island, San Juan Island 122°59'48” 48°32/15” D. frondosus, D. rufus, D. albus, D. diversicolor
Cantilever Pier, Friday Harbor 123°00'18” 48°32'48” D. frondosus, D. diversicolor
Laboratories, San Juan Island
Clarke Rock, Nanaimo, B. C. 123°56/30” 49° 13/30” D. iris
Collins Cove, San Juan Island 123°00’18” 48°33" D. albus, D. diversicolor
Departure Bay, Nanaimo, B. C. 1239577241" 49° 12’45” D. iris, D. rufus
Eagle Point, San Juan Island 123°02’42” 48°27'30” D. dalli, D. diversicolor
East Sound, Orcas Island 122°58’30” 48°30! D. iris
Edwards Reef, San Juan Island 123°07/36” 48°29/48”” D. frondosus, D. diversicolor, D. albus, D. subramosus
Harney Channel, Lopez Island 122°50’30” 48°34’ D. iris, D. dalli
Lonesome Cove, San Juan Island 123°07’ 48°37'30” D. dalli, D. diversicolor, D. subramosus, D. albus
Low Island, San Juan Island 123°10’12” 48°39'42” D. subramosus
Mitchell Bay, San Juan Island 123°10'24” 48°34/12”” D. frondosus
Peavine Pass, Blakely Island 122°48/36” 48°35/24”” D. diversicolor, D. subramosus
Point Caution, San Juan Island 123°00'48” 48°33'42” D. diversicolor
Breakwater, Victoria f 123°23’36” 48°24'48” D. frondosus, D. dalli, D. diversicolor
Turn Rock, San Juan Island 122°57'48” 48°32/06” D. dalli
West Sound, Orcas Island 122°58’ 48°37’ D. iris
Vol. 12; No. 4
THE VELIGER
Page 477
APPENDIX II
Synonymy of Morphological Terms of Other Authors
In the widely scattered nudibranch literature, numerous
terms are often applied to the same structure or, converse-
ly, the same term may be applied to entirely different
structures. In an effort to dispel some of this confusion,
I have compiled a list of some of the more common terms
that other authors have given to structures described in
the text. Normally, the term I have used is the one used
most frequently in the literature at present. If the term
is new, it is defined in the text.
The synonym is listed under the appropriate heading,
followed by the name of the author who used it in this
sense. Finally, the term used in this paper to which it
is synonymous, is given.
Term Author Presently used Term
Body
cardiac elevation MacFar.ianp, 1966 cardiac prominence
Cerata
branchiae ALpER & Hancock, 1845 cerata
VERRILL, 1870
branchial processes Cooper, 1863 cerata
back papillae Bercu, 1894 cerata
gills Bercy, 1894 cerata
dorsal papillae Bercu, 1879 cerata
dorso-lateral processes © MacFarianp, 1966 cerata
Rhinophore and Clavus
tentacle ALpER & Hancock, 1845 clavus
tentacle sheath
branches
tentacles
club
arborescent papillae
Head, Lips and Veil
papillae
frontal processes
frontal sail
arbusculi
velum
Jaws
chewing process
mandible
Radula
tongue
hook of lateral tooth
Reproductive System
ovarium
vesicula seminalis
spermatotheca
vaginal duct (proximal
portion only)
spermatheca
VERRILL, 1870
ALDER & Hancock, 1845
VERRILL, 1870
O’DonocHuuE, 1921
ALDER & Hancock, 1845
Cooper, 1863
Bercu, 1879
OpHNe_R, 1939
MacFaruanp, 1966
Bercu, 1894
Bercu, 1894
Bercu, 1894
Bercu, 1879
O’DonocHuE, 1921
MacFarianp, 1966
Bercu, 1894
MacFarianp, 1966
ALDER & Hancock, 1845
rhinophore sheath
and stalk
crown papillae
whole rhinophore
clavus.
lateral papillae
lip papillae
veil papillae
veil
veil papillae
lateral papillae
masticatory process
jaw (mandible might
be considered a better
term; my choice was
arbitrary)
radula
Other authors use the term tongue, but
usually when referring to the odontophore
which supports the radula
MacFar.anp, 1966
ALDER & Hancock, 1845
Opune_r, 1936
MacFarianp, 1966
MacFar.anp, 1966
GuIsELIN, 1965
cusp of lateral tooth
hermaphrodite gland
seminal receptacle
seminal receptacle
insemination duct
bursa copulatrix
————————————————————
Page 478
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Marcus, ERNST
1961. Opisthobranch mollusks from California. The Veli-
ger 3 (Supplement, pt. 1): 1-85; plts. 1 - 10.
(1 February 1961)
Mayr, ERNsT
1963. Animal species and evolution.
Harvard Univ. Press; 797 pp.; illust.
Meyer, Heinrich Aucust & Kart Mostus
1865. Fauna der Kieler Bucht. I. Die Hinterkiemer oder
Opisthobranchia. viii + xxx + 88 pp.; 26 plts. Leipzig,
Wilhelm Engelmann, Verlag.
Mitier, MicHaEL CHARLES
1961. Distribution and food of the nudibranchiate Mollusca
of the South of the Isle of Man. Journ. Anim. Ecol. 30:
95 - 116
1962. Annual cycles of some Manx nudibranches with a dis-
cussion of the problem of migration. Journ. Anim. Ecol.
31: 545 - 569
Morton, JoHN Epwarp
1958. Molluscs.
23 text figs.
Mutter, Otro FREDERIK
1776. _Zoologiae Danicae Prodromus.
Opuner, Nits HJALMaR
1926. Nudibranchs and lamellarids from the Trondhjem
Fjord. K. Norske Vidensk. Selsk. Skr. pp. 1 - 36
1934. The Nudibranchiata. In British Antarctic (“Terra
Nova’) Expedition, 1910. Nat. Hist. Rep. Zool. 7 (5):
229-310; plts. 1-3. London.
1936. | Nudibranchia Dendronotacea. A revision of the System.
Meélanges Paul Pelseneer. Mém. Mus. Roy. d’Hist. Nat.
de Belgique, Ser. II, Fasc. 3: 1057-1128; 1 plt.; text figs. 1-47.
1939. | Opisthobranchiate mollusca from the western and north-
erm coasts of Norway. K. Norske Vidensk. Selsk. Skr. no.
1: 1-92; 59 text figs.
Cambridge, Mass.
Hutchison Univ. Libr., London. 232 pp.;
p- 229
Vol. 12; No. 4
O'DonocHuE, CHARLES HENRY
1921. Nudibranchiate Mollusca from the Vancouver Island
region. Trans. Roy. Canad. Inst. 13 (1): 147 - 209
1922. Notes on the nudibranchiate Mollusca from the Van-
couver Island region. III. Records of species and distribution.
Trans. Roy. Canad. Inst. 14 (1): 145 - 167; plts. 5, 6
1924. Notes on the nudibranchiate mollusca from the Vancouver
Island region. IV. Additional species and records. Trans.
Roy. Canad. Inst. 15 (1): 1-33; plts. 1, 2
Plymouth Marine Invertebrate Fauna
1904. Journ. Marine Biol. Assoc. U.K. 7: 155 - 298
Pruvot-Fot, ALICE
1954. Mollusques opisthobranches. In Faune de France, 58:
460 pp.; 1 plt.; 173 text figs.; Paris, Paul Lechevalier.
RosiLiiarpD, Gorpvon A.
1969. A method of color preservation in opisthobranch mol-
luscs. The Veliger 11 (3) : 289 - 291 (1 January 1969)
Smiru, Atityn G., & MacKenzie Gorpon, Jr.
1948. The marine mollusks and brachiopods of Monterey
Bay, California, and vicinity. Proc. Calif. Acad. Sci.,
4th. Ser., 26 (8): 147 - 245; plts. 3-4; textfigs. 1-4
STAsEK, CHARLES ROBERT
1967. Autotomy in the Mollusca.
Sci. 61: 1 - 44
SWENNEN, C.
Occ. Pap. Calif. Acad.
1961. . Data on distribution, reproduction, and ecology of the
nudibranchiate Mollusca occurring in the Netherlands.
Netherl. Journ. Sea Res. 1 (1-2): 191-240
THIELE, JOHANNES
1929. Handbuch der systematischen Weichtierkunde. (Jena,
Gustav Fischer, 1929 - 1935) ; 1154 pp.; 893 text figs.
TuHompson, THomas E.
1960a. On a disputed feature of the anatomy of the nudibranch
Dendronotus frondosus ASCANIUS. Proc. Malac. Soc. Lon-
don 34 (1): 24-26
1960b. Defensive acid secretion in marine gastropods.
Journ. Marine Biol. Assoc. U.K. 39: 115 - 122
THE VELIGER
Page 479
1961. The structure and mode of functioning of the reproduc-
tive organs of Tritonia hombergi. Quart. Journ. Micr. Sci.
102: 1 - 14
1964. Grazing and the life cycles of British nudibranchs.
Brit. Ecol. Soc. Symp. 4: Grazing in terrestrial and marine
environments, ed. D. J.Crisp. Blackwell, Oxford: 275 - 297
VERRILL, ADDISON EMERY
1870. Contributions to Zoology from the Museum of Yale
College. No. 8 — Descriptions of some New England Nudi-
branchiata. Amer. Journ. Sci. Arts 50: 405 - 406
1879. Notice of some recent additions to the marine Inverte-
brata of the northeastern coast of America, with descriptions of
new genera and species and critical remarks on others.
Proc. U.S. Nat. Mus. 2: 165 - 205
1880. Notice of some recent additions to the marine Inverte-
brata of the northeastern coast of America, with descriptions of
new genera and species and critical remarks on others. Part II.
Mollusca with notes on Annelida, Echinodermata, etc. collected
by the U.S. Fish Commission. Proc. U.S. Nat. Mus. 3:
356 - 405
1882. Catalogue of marine Mollusca added to the fauna of
the New England region during the past ten years. Trans.
Conn. Acad. Arts Sci. 5: 447 - 587
VoLopcHENKO, N. I.
1955. Subclass Opisthobranchia, pp. 247 - 25 ; plt. 48.
In: Atlas of the invertebrates of the far eastern seas of the
USSR, E. N. Paviovsxu [ed.]. Transl. by A. Marcapo, 1966.
Wa ter, RicHarD A. & WILL1AM N. ESCHMEYER
1965. A method for preserving color in biological specimens.
Bioscience 15 (5): 361
WATERS, VIRGINIA
1966. Feeding ecology and other aspects of the natural history
of the nudibranch, Eubranchus olivaceus. M. S. Thesis,
Univ. Washington, Seattle, Wash.
Wo ttTerR, HELMA
1967. __ Beitrage zur Biologie, Histologie und Sinnesphysiologie
(insbesondere der Chemorezeption) einiger Nudibranchier
[Mollusca, Opisthobranchia] der Nordsee. Zeitschr. Morph.
Okol. Tiere 60: 275 - 337
Page 480
THE VELIGER
Vol. 12; No. 4
Notes on the Egg Capsules and Larval Development of
Conus purpurascens BRODERIP
BY
JAMES NYBAKKEN
Moss Landing Marine Laboratories, Box 223, Moss Landing, California 95039
(10 Text figures)
THE EGG CAPSULES of 11 species of Indian Ocean Conus
and the larval development of 4 species have been report-
ed by Konn (1961a). Korn (1961b) has also described the
egg capsules of 9 Hawaiian Conus species and the larval
development of 4 of these. However, no descriptions of
breeding season, egg capsules or larval development of
the West American Conus species are known to this
author. This note describes the egg capsules of Conus
purpurascens Broverip, 1833, and some features of the
early larval development.
A single large Conus purpurascens was found on the
underside of a rock 15 inches in diameter at a depth
of 2 feet just outside San Carlos Bay near Guaymas,
Sonora, Mexico on July 28, 1966. The animal was in
the process of ovipositing when discovered. About 150
egg capsules were attached to the underside of the rock
at that time. It is not known if all were deposited by this
single female, but no other individuals of C’. purpurascens
were found near the rock. However, 4 other specimens of
this same species were collected in the same general area.
The egg capsules were typical of Conus egg capsules
as described by Koun (1961a, b) and were attached in
irregular rows of 10-20 cases (Figure 1). Some cases
were deposited upon the top of others. The capsules varied
in height from 9.6 to 15.0 mm and from 9.3 to 11.4mm
in width. Capsules were translucent white with the en-
closed eggs readily visible inside.
Figure 1
Left: a single egg capsule of Conus purpurascens viewed from the
side; right: a group of three Conus purpurascens egg capsules
viewed from the front
Several capsules were collected and taken to the Moss
Landing Marine Laboratories where they were kept in
finger bowls of sea water which was changed every 3 days.
The number of eggs per capsule ranged from 5400 to
5900 which is considerably higher than the 1650 re-
ported by Koun (1961b) for Conus catus BRUGUIERE,
1792, another fish eating species. The number falls more
within the range reported in capsules of the vermivorous
species.
Unfortunately, due to the lack of optical equipment in
Mexico and the delay in transit back to Moss Landing,
the collected egg capsules were not examined to deter-
mine the stage of development until August 5, 1966.
Hence it is not possible to say what the stage of develop-
ment was when the capsules were collected or what
happened up until August 5".
On August 5" a study was begun of 3 selected capsules
to trace development. When examined at that time the
capsules contained live embryos that were primarily in
a trochophore-like stage of development (Figures 2, 3).
0.1 mm
Figure 2
Early trochophore-like larva of Conus purpurascens
Figure 3
Trochophore-like larva of Conus purpurascens
The size of 9 measured trochophores averaged 0.184 mm
in length. The trochophores were very irregular in shape
which corroborates Kouwn (1961a).
Vol. 12; No. 4
THE VELIGER
Page 481
\
On August 10" the capsules still had many of these
trochophore-like stages, but many of larvae appeared to
be developing velar lobes (Figures 4, 5) and by August
11" most were in early veliger stage (Figure 6). That
the stage of development was not consistent even within
Figure 4
Early veliger stage of Conus purpurascens
Figure 5
Early veliger stage of Conus purpurascens
VL = velar lobes
single capsules became apparent by August 14". At that
time most of the larvae were in early or late veliger
stage (Figures 6, 7), but it was possible to find trocho-
phores and even what appeared like early cleavage stages
Sh
Figure 6
Early veliger stage of Conus purpurascens
Figure 7
Veliger stage of Conus purpurascens
VL = velar lobes VM = visceral mass SH = shell
in the same capsule. Such a situation was not reported by
Koun (196la, b) and may have been due to the un-
natural conditions to which the capsules had been sub-
jected since collection and to the fluctuating temperatures
of the room in which they were kept.
0.1 mm
Figure 8
Late veliger stage of Conus purpurascens
f = foot Ps = pigment spots
Sh = shell
Op = Operculum
On August 20" most of the larvae were in a well
developed veliger stage in which they remained for the
next several days (Figure 8). The size of the veliger shells
averaged 0.2011 mm [16 shells measured] and the average
total length of 11 veligers was 0.237mm. No further
development was observed, and by August 26" most veli-
gers were dead in the capsules. Hatching was not ob-
served in any of the cases. It is not known if hatching
occurs in nature in the veliger stage.
Figure 9
Late veliger stage of Conus purpurascens
h = heart VL = velar lobes
Sh = shell
VM = visceral mass
Figure 10
Veliger shell of Conus purpurascens
LITERATURE CITED
Koun, ALAN JAcoBs
1961a. Spawning behavior, egg masses, and larval development
in Conus from the Indian Ocean. Bull. Bingham Ocean. Coll.
17 (4): 51 pp.
1961b. Studies on spawning behavior, egg masses, and larval
development in the gastropod genus Conus. Part I. Observa-
tions on nine species in Hawaii. Pacif. Sci. 15 (2) :163 - 179
Page 482
THE VELIGER
Vol. 12; No. 4
A Supplement to the Annotated List of Opisthobranchs
from San Luis Obispo County, California
RICHARD A. ROLLER
1127 Seaward Street, San Luis Obispo, California 93401
SINCE COMPILATION OF the original list of opisthobranchs,
the continued collection of specimens and additional data
from San Luis Obispo County require a supplement to
that list. Also, two more inferred species for the county,
and an omission from the original “Literature Cited” sec-
tion should be added at this time.
I would like to acknowledge the kind assistance of Mr.
Gary R. McDonald and Mr. Steven J. Long for furnishing
collecting data and specimens.
LIST or COLLECTED SPECIES
Acanthodoris hudsont MacFarLanp, 1905
[see McDonatp, 1970]
Rare.
Morro Bay, Hazard Canyon, Shell Beach.
Vancouver Island to Gaviota, Santa Barbara County.
Acteocina culcitella (Goutp, 1853)
Rare.
Morro Bay.
Kodiak Island to San Diego.
Acteocina inculta (Gout p, 1855)
Common.
Morro Bay.
Monterey Bay to Gulf of California.
Acteon punctocaelatus (CARPENTER, 1864)
Rare.
Cayucos, Morro Bay.
Southeastern Alaska to Magdalena Bay, Baja California.
Aglaja diomedea (Brrcu, 1894)
Common.
Morro Bay.
Kodiak Island to Morro Bay (Kodiak Island to Elkhorn
Slough).
Aplysia vaccaria WINKLER, 1955
Rare.
Morro Bay.
Morro Bay to Gulf of California (Santa Barbara County
to Gulf of California).
Chromodoris macfarlandi (CocKERELL, 1902)
Rare. Subtidal to 20 feet.
Shell Beach.
Monterey Bay to Ensenada, Baja California, Mexico.
Cumanotus beaumonti (Eutot, 1908)
Rare. Found on Tubularia sp. on floating docks.
Morro Bay.
Friday Harbor, Washington to San Diego; Norway, Great
Britain.
Dendronotus iris Cooper, 1863
Rare. Subtidal to 20 feet.
Morro Bay.
Vancouver Island to Coronados Islands.
Elysia hedgpethi Marcus, 1961
Rare.
Shell Beach, Morro Bay.
San Juan Island, Washington, to La Jolla, California;
Bahia de Los Angeles, Baja California.
Onchidoris bilamellata (Linnaeus, 1767)
[see McDona tp, 1970]
Frequent. On floating docks, buoys, and subtidal to 20 feet.
Morro Bay.
Alaska to Morro Bay; circumboreal.
Onchidoris muricata (MuLuER, 1776)
[listed as Onchidoris sp. in ROLLER « Lone, 1969]
Rare.
Hazard Canyon, Lion Rock.
Friday Harbor, Washington to Lion Rock, San Luis Obis-
po County; northern Europe.
Vol. 12; No. 4
Placida dendritica (ALDER & Hancock, 1843)
[see Lone, 1969]
Rare. On Bryopsis corticulans SETCHELL.
Shell Beach, Morro Bay.
Fort Barry Boat Harbor, Marin County, California to
Shell Beach; North Carolina, coast of New England;
Mediterranean Sea and eastern Atlantic; Japan.
Trinchesia fulgens (MaAcFarLanp, 1966)
[see Lone, 1969]
Rare.
Shell Beach, Hazard Canyon.
Duxbury Reef, Bolinas Bay to Shell Beach.
Trinchesia virens (MAcFartanp, 1966)
[see Lone, 1969]
Rare.
Shell Beach.
Monterey Bay to Shell Beach.
LIST or INFERRED SPECIES
Gastropteron pacificum BrercH, 1894
[see BerTscH, 1969]
Aleutian Islands to Galapagos Islands.
Pleurobranchaea californica MAcFaRLAND, 1966
[see Cutvers, 1967]
Mouth of Klamath River, California, to San Diego.
LITERATURE CITED
ALDER, JosHUA & ALBANY Hancock
1843. Notice of a British species of Calliopaea d’Orb. and of
four new species of Eolis, with observations on the development
and structure of the nudibranchiate mollusca. Ann. Mag.
Nat. Hist. 12: 233 - 238 [not seen]
Bercy, Lupwic SopHus RuDOLPH
1894. Die Opisthobranchien. Bull. Mus. Comp. Zool.
Harvard 25 (10): 125 - 233; plts. 1-12
THE VELIGER
Page 483
BertscH, Hans
1969. A note on the range of Gastropteron pacificum. The
Veliger 11 (4) : 431 - 433; 1 fig. (1 April 1969)
CARPENTER, PHILIP PEARSALL
1864. | Supplementary report on the present state of our know-
ledge with regard to the Mollusca of the west coast of North
America. Rept. Brit. Assoc. Adv. Sci. for 1863: 517 - 686
(August 1864)
[not seen]
Cuivers, Dustin DALE
1967. Observations on Pleurobranchaea californica MacFar-
LAND, 1966 (Opisthobranchia, Notaspidea) . Proc. Calif.
Acad. Sci. 32 (17): 515-521; figs. 1-4 (22 Nov. 1967)
CocKERELL, THEODORE Dru ALLISON
1902. Three new species of Chromodoris.
19-21
Cooper, JaMES GRAHAM
1862. Some genera and species of California Mollusca.
Proc. Calif. Acad. Nat. Sci. 2: 202 - 207
Franz, Davp R.
1967. On the taxonomy and biology of the dorid nudibranch
Doridella obscura. The Nautilus 80 (3): 73-79; 1 fig.
LinnaEus, Caro.us
1767. Systema naturae per regna tria naturae ... editio
duodecima, reformata 1 [Regnum animale] (2): 533 - 1327.
Stockholm (Laurentii Salvii) °
[not seen]
Nautilus 16:
Lone, STEVEN J.
1969. Records of Trinchesia virens, Trinchesia fulgens, and
Placida dendritica from San Luis Obispo County, California.
The Tabulata 2 (4): 9-12; 2 figs. (1 October 1969)
MacFar.anpb, FRANK MAcE
1905. A preliminary account of the Dorididae of Monterey
Bay, California. Proc. Biol. Soc. Washington, 18: 35 - 54
1966. Studies of opisthobranchiate mollusks of the Pacific
Coast of North America. Mem. Calif. Acad. Sci. 6: xvi +
546 pp.; 72 plts. (8 April 1966)
Marcus, ERNST
1961. Opisthobranch mollusks from California. The
Veliger 3 (Supplement, pt. I): 1-85; plts. 1-10. (Feb. 1, 1961)
McDona tp, Gary R.
1970. Range extensions for Acanthodoris hudsont MacFar-
LAND, 1905, and Onchidoris bilamellata (LinnaEus, 1767).
The Veliger 12 (3) : 375 (1 January 1970)
Rotter, RicHarp A. & STEVEN J. Lonc
1969. An annotated list of opisthobranchs from San Luis Obis-
po County, California. The Veliger 11 (4): 424-430; 1
map (1 April 1969)
Page 484
NOTES & NEWS
Concerning the Validity
of the New Species of Paravitrea Proposed
by BRANSON & BATCH, 1970
BY
LANDON T. ROSS
AND
LAWRENCE G. ABELE
Department of Biological Science
Florida State University, Tallahassee, Florida 32306
IN A RECENT publication, Branson & Batcu (1970)
inadvertently introduced into the literature two unavail-
able and invalid scientific names. They noted that 52
species of land and freshwater gastropods were collected
from an area in northern Kentucky, and that two of
these species were “apparently new” (p. 338). The au-
thors stated that these species were to be described, but
not named (p. 338). The two species in question were
listed as “Paravitrea species a” and “Paravitrea species
b” (p. 341). “Paravitrea species a” was described under
the heading “Paravitrea new species? a” (p. 342) and
“Paravitrea species b” under the heading “Paravitrea
new species? b” (p. 343). Unfortunately, in figure 4 of
the same article (p. 347) five species of Paravitrea (the
same number of species in that genus as recorded in the
body of the report including species “a” and “b”) are
listed by name. Two of these names are new and may be
easily correlated with Paravitrea species “a” and “‘b’’ by
means of the listed locality data. “Paravitrea species a”
is P lafuzi BRANSON & Batcu, 1970, and Paravitrea spe-
cies b” is P sceada BRANSON & Batcu, 1970.
It is evident that the authors did not wish these species
to be named, possibly because of an insufficient number
of specimens, and that provisional names accidentally
crept into the manuscript. However, since “‘new species?”
was included in the description of both species, the names
may be considered to have been proposed conditionally
under the provisions of Article 15 of the International
Code of Zoological Nomenclature (1964). These names,
THE VELIGER
Vol. 12; No. 4
Paravitrea lafuzi BRANSON & Batcu, 1970, and Paravit-
rea sceada BRANSON & Batcu, 1970, are not available
taxonomically. If the two species are to be eventually
renamed, two new names must be proposed.
LITERATURE CITED
BRANSON, BRANLEY ALLAN & Donatp L. Batcu
1970. An ecological study of valley-forest gastropods in a
mixed mesophytic situation in northern Kentucky. The
Veliger 12 (3): 333 - 350; 7 text figs. (1 January 1970)
Stott, Norman Rupo.pH, ef al.
1964. International Code of Zoological Nomenclature adopted
by the XV International Congress of Zoology, ed. 2.
London (Internat. Trust f. Zool. Nomencl.) pp. i-xvii+
1-176; 5 appendices & Glossary
A. M. U.
The 36" annual meeting of the American Malacological
Union, Inc. will be held from July 16 to July 20, 1970 in
Key West, Florida. This coincides with the minus tides to
increase the collecting opportunities and pre-dates the
hurricane season. Special symposia on biological system-
atics of marine bivalves and gastropods, commercial shell
fish, and introduced mollusks, are being organized. Several
features of interest to the amateur will be announced
later. Information on program and housing can be ob-
tained from Dr. Alan Solem of the Field Museum of
Natural History, Roosevelt Road at Lake Shore Drive,
Chicago, Illinois 60605.
W. S. M.
The Third Annual Meeting of the Western Society of
Malacologists is scheduled to be held at Stanford Uni-
versity, June 24 through June 27, 1970. Besides general
papers, the program that is being planned includes a
symposium on Advances in Molluscan Systematics: a
Survey of New Theory and Practice. Dr. Eugene Coan,
who is organizing the symposium, reports that papers
have been promised in the fields of microstructure of
shells, radiography, biochemistry, and others.
Housing will be in university dormitories, with cafe-
teria meals, probably in the dormitory area.
Exhibit Chairman is Twila Bratcher; Hospitality Chair-
man, Mary D’Aiuto.
Myra Keen, President, 1970
Vol. 12; No. 4
THE VELIGER
Page 485
International Commission on Zoological Nomenclature
Opinion 885
(October 1969)
The name of the type of Mitra is interpreted to be Voluta
mitra LinNAEUS, 1758, not Voluta episcopalis
Linnaeus, 1758.
Opinion 886
(October 1969)
Ocenebra Gray, 1847 put on the Official List of Generic
Names.
Tritonalia FLEMING put on the Official Index of Rejected
Generic Names.
The type of Purpura is declared to be P persica, which
fixes the concept of the genus in its present sense.
The type of Muricanthus is fixed as Murex radix.
The family name Thaididae is given precedence over the
earlier Purpuridae.
Two specific names by Perry, 1810, are suppressed —
pictum and dentex.
Correction!
Our readers will, no doubt, have noticed that the explan-
ations for Figures 5 and 6 on Plate 52 of our January
issue are reversed. We are sorry that we did not catch
this error.
Important N otices
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In the face of continuous rises in the costs of printing
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is the stated aim of the Society to disseminate new infor-
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At a Regular Membership meeting of the Society i in No-
vember 1968 a policy was adopted which, it is hoped,
will assist in building up the Endowment Fund of the
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An issue of the journal will be designated as a Memorial
Issue in honor of a person from whose estate the sum of
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will be dedicated to the memory of the decedent.
Vol. 12; No. 4
CALIFORNIA
MaA.acozoo.ocicat Society, Inc.
is a non-profit educational corporation (Articles of In-
corporation No. 463389 were filed January 6, 1964 in
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tiate their respective tax deductions.
As has been announced in several previous issues of
our journal, changed postal rules affecting second class
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customary reminders in our January issue, nor can we
attach flyers to our journal. We have stated, however, that
we Shall print a statement in the NOTES & NEWS sec-
tion when renewal time has arrived.
Therefore we now make the announcement that re-
newals of memberships are due. The Regular Membership
at its meeting on November 10, 1969, decided unani-
mously to hold dues and subscription rates at their present
level in spite of the continued inflationary pressures.
Statements and invoices are now in the mail and may
arrive even before the current number. Prompt payment
of the dues, to reach the Manager before April 15, 1970,
will assist in keeping expenses - and, consequently, the
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Jean M. Cate, Manager.
THE VELIGER
Page 487
BOOKS, PERIODICALS, PAMPHLETS
Olive Shells of the World
by Row1anp F ZeicLer & Humsert C. Porreca. Ro-
chester, N. Y.: 96 pp., 13 colored plts., 6 unnumbered plts.
price, $12.95. The book is being distributed by Richard E.
Petit, P.O. Box 133, Ocean Drive Beach, South Carolina,
29582.
This is a well arranged and beautifully bound book.
The introduction discusses shell characters, anatomical
structures, feeding and breeding habits of the snails, and
how to go about collecting them. Classification is given
briefer treatment, though the lists of available subgeneric
names and the lists of synonyms under each species head-
ing let the collector know of their existence. All currently
recognized species are illustrated at natural size in color,
often with more than one figure to show variation. Many
of the named color varieties are figured. In the text these
are cited as “forma,” and their infrasubspecific status is
made clear. However, in the plate captions the names are
treated as trinomials, as if they were true subspecies. This
may be a source of confusion. There is a brief but ade-
quate bibliography and a good index; also a useful two-
page glossary. The color plates are grouped under three
geographic areas -— Atlantic, East Pacific, and Indo-
Pacific — and the species are listed alphabetically (not
systematically) for each area. In the introduction the
species names are grouped under subgeneric categories,
the bases of which are not cited.
Color reproduction is good, and the plates are large
enough (8X11 inches) so that a sizable fraction of the
species in each area can be lined up for direct compar-
ison. Obviously the book has been put together with care
and thought. It seems to be accurate -— not even a
typographical error was detected in a first-time-through
reading. Thus it can be recommended to anyone wishing
a modern reference work on this colorful but somewhat
neglected family of marine gastropods.
MK
Marines Pliozan und Pleistozan in Nord- und Mittel-
Chile unter besonderer Beriicksichtigung
der Entwicklung der Mollusken-Faunen.
by Dretricu Herms. Zitteliana: Abhandlungen der Bay-
erischen Staatssammlung fiir Palaontologie und historische
Geologie, no. 2, pp. 1-159; plts. 1-18. September 1,
1969 (Munich, Germany).
Page 488
THE VELIGER
Vol. 12; No. 4
This is a detailed study of the sedimentary history of
late Cenozoic marine strata from the Peruvian border
south to Coquimbo, Chile. Successive faunal changes are
carefully documented, and the use of paleo-ecological
analyses enables the recognition of several faunal assem-
blages. The text is in German, with summaries in English
and Spanish. Three new taxa are described: Patella fuen-
zalidai, Anomia atacamensis, and Protothaca antiqua ton-
goyensis. The 18 plates are of excellent photographic
quality. The work also has an extensive bibliography. It
is a major contribution in its field.
Availability and price are not indicated on the paper
cover; presumably copies would be obtainable from the
Bayerische Staatssammlung fir Palaontologie und histo-
rische Geologie, 8 Munchen 2, Richard-Wagner Strafe
10/11, Germany.
MK
Die europaischen Meeresmuscheln (Bivalvia)
vom Eismeer bis Kapverden, Mittelmeer
und Schwarzes Meer
by Dr. Frrrz Norpsieck. 1969. xiii+ 256 pp.; 900
line drawings on 26 plates; 7 color pictures on 2 plates.
Hardcover, 48.- German Reichsmarks; Gustav Fischer,
publ., Stuttgart, West Germany.
In our Volume 11, on page 444, we reviewed what now
turns out to have been the first volume on mollusks from
the area delimited in the title of the book. The same
author has brought the same painstaking attention to
the multitudinous details again. One important addition
is a list of abbreviations used in this volume. This will
now make the book even more valuable to the worker
whose knowledge of the German language is not of the
very best.
The general plan for this book is exactly the same as
that adopted in the first volume. We think that it is
indispensable for any worker concerned with European
marine bivalves and it certainly is a welcome tool for
all curators of collections.
The last 3 pages of this book contain additions to the
previous volume.
With a bit of nostalgia, this reviewer remembers with
great pleasure the many fine books, lavishly illustrated,
for which this publishing house has been justly famous.
It seems, considering modern trends in publishing and in
printing methods, that the publisher continues to uphold
the high standards set by the original founder of the firm.
RS
West Coast Chitons
A Collector’s Guide to West Coast Chitons
by GLENN & Laura BurcHarnt. Special publication no.
4, San Francisco Aquarium Society, Inc. 45 pp.; 80 color
illustr. on 4 plts., 6 figs. on 1 halftone plt. and 1 text fig.,
price $4.95; available from the authors. Nov. 1969.
An extraordinarily fine color photograph of the rare
Mopalia lowe: Prtspry, 1918, adorns the front cover of
this spiral-bound book, the size of which is 84 by 11
inches. In the introduction the authors stress the existent
lack of appropriate literature by which the collector can
identify the chitons which are fairly abundant on the
west coast of North America. To erase this lack is the pur-
pose of this informally written work. A chapter on “Chi-
tons in General” is followed by one on “Collecting and
Preparing Chitons.” The main body, which follows, enu-
merates, in alphabetical order, all the species the authors
have in their collections. The information given includes
the currently accepted valid scientific name of the species,
the range, the type locality. a brief description of the
valves and animal and, where appropriate, the syno-
nymy. A glossary of terms applicable to chitons, copied
from ARNOLD’s glossary of 1001 terms used in conchology
(Supplement to vol. 7 of the Veliger) precedes the final
3 pages of “Photo Descriptions & Credits.”
The authors have omitted the dates of original des-
criptions, as well as the parentheses in those cases where
species have been assigned to genera different from the
original assignment.
This work will be welcomed by all the enthusiastic
collectors of this very interesting group of mollusks, repre-
sented in such a varied and large number of species.
RS
Marine Shells of Southern California
by James H. McLean. Science Series 24, Zoology No. 11,
Los Angeles County Museum of Natural History, Octo-
ber, 1969 104 pp.; several 100 photographs reproduced
as 54 figures. Price $2.00.
This small book contains a wealth of well-organized and
well presented information. Taxonomically it is up to
date. The photographs of every species treated are of
high quality. The modest price of the book should make
it possible for every serious student of the molluscan fauna
of California to add it to his library.
RS
EDITORIAL BOARD
Dr. Donatp P. Azssott, Professor of Biology
Hopkins Marine Station of Stanford University
Dr. Jerry DonouueE, Professor of Chemistry
University of Pennsylvania, Philadelphia, and
Research Associate in the Allan Hancock Foundation
University of Southern California, Los Angeles
Dr. J. Wyatr Duruam, Professor of Paleontology
University of California, Berkeley
Dr. E. W. Facer, Professor of Biology
Scripps Institution of Oceanography, La Jolla
University of California at San Diego
Dr. Caper Hanp, Professor of Zoology and
Director, Bodega Marine Laboratory
University of California, Berkeley
Dr. G Datias Hanna, Curator
Department of Geology
California Academy of Sciences, San Francisco
Dr. Jorn W. HepcpetH, Resident Director
Marine Science Laboratory, Oregon State University
Newport, Oregon
Dr. Leo G. HERTLEIN,
Curator of Invertebrate Paleontology
California Academy of Sciences, San Francisco
EDITOR-IN-CHIEF
Dr. Rupotr STouiER, Research Zoologist
University of California, Berkeley
Dr. A. Myra KEEN, Professor of Paleontology and
Curator of Malacology
Stanford University, Stanford, California
Dr. Victor Loosanorr, Professor of Marine Biology
Pacific Marine Station of the University of the Pacific
Dr. Joun McGowan, Associate Professor of
Oceanography
Scripps Institution of Oceanography, La Jolla
University of California at San Diego
Dr. Frank A. Pitetxa, Professor of Zoology
University of California, Berkeley
Mr. Attyn G. Situ, Associate Curator
Department of Invertebrate Zoology
California Academy of Sciences, San Francisco
Dr. Ratpu I. Smiru, Professor of Zoology
University of California, Berkeley
Dr. Cuartes R. STASEK, Associate Professor
of Zoology
Florida State University, Tallahassee, Florida
Dr. Donatp M. Witson, Professor of Biology
Department of Biological Sciences
Stanford University, Stanford, California
ASSOCIATE EDITOR
Mrs. JEAN M. Cate
Los Angeles, California
THE VELIGER is open to original papers pertaining to any problem concerned
with mollusks.
This is meant to make facilities available for publication of original articles
from a wide field of endeavor. Papers dealing with anatomical, cytological, distri-
butional, ecological, histological, morphological, physiological, taxonomic, etc.,
aspects of marine, freshwater or terrestrial mollusks from any region, will be
considered. Even topics only indirectly concerned with mollusks may be acceptable.
It is the editorial policy to preserve the individualistic writing style of the
author; therefore any editorial changes in a manuscript will be submitted to the
author for his approval, before going to press.
Short articles containing descriptions of new species or other taxa will be given
preferential treatment in the speed of publication provided that arrangements
have been made by the author for depositing the holotype with a recognized
public Museum. Museum numbers of the type specimens must be included in the
manuscript. Type localities must be defined as accurately as possible, with geo-
graphical longitudes and latitudes added.
Short original papers, not exceeding 500 words, may be published in the column
“NOTES and NEWS’; in this column will also appear notices of meetings of
regional, national and international malacological organizations, such as A.M. U.,
U.M.E., W.S.M., etc., as well as news items which are deemed of interest to
our Members and subscribers in general. Articles on “METHODS and TECH-
NIQUES” will be considered for publication in another column, provided that
the information is complete and techniques and methods are capable of duplication
by anyone carefully following the description given. Such articles should be mainly
original and deal with collecting, preparing, maintaining, studying, photographing,
etc., of mollusks or other invertebrates. A third column, entitled “INFORMA-
TION DESK,” will contain articles dealing with any problem pertaining to
collecting, identifying, etc., in short, problems encountered by our readers. In
contrast to other contributions, articles in this column do not necessarily contain
new and original materials. Questions to the editor, which can be answered in this
column, are invited. The column “BOOKS, PERIODICALS, and PAMPHLETS”
will attempt to bring reviews of new publications to the attention of our readers.
Also, new timely articles may be listed by title only, if this is deemed expedient.
Manuscripts should be typed in final form on a high grade white paper, not
exceeding 81/,” by 11”, at least double spaced and accompanied by a clear carbon
or photo copy. A pamphlet with detailed suggestions for preparing manuscripts
intended for publication in THE VELIGER is available to authors upon request.
A self-addressed envelope, sufficiently large to accommodate the pamphlet (which
measures 514,” by 814""), with double first class postage, should be sent with the
request to the Editor.
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